Trees, Crops and Soil Fertility: Concepts and Research Methods

TREES, CROPS AND SOIL FERTILITY CONCEPTS AND RESEARCH METHODS Trees, Crops and Soil Fertility Concepts and Research M...

Author: G Schroth | F L Sinclair

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TREES, CROPS AND SOIL FERTILITY CONCEPTS AND RESEARCH METHODS

Trees, Crops and Soil Fertility Concepts and Research Methods

Edited by

G. Schroth Biological Dynamics of Forest Fragments Project National Institute for Research in the Amazon Manaus Brazil

and

F.L. Sinclair School of Agricultural and Forest Sciences University of Wales Bangor UK

CABI Publishing

CABI Publishing is a division of CAB International CABI Publishing CAB International Wallingford Oxon OX10 8DE UK Tel: +44 (0)1491 832111 Fax: +44 (0)1491 833508 E-mail: [email protected] Website: www.cabi-publishing.org

CABI Publishing 44 Brattle Street 4th Floor Cambridge, MA 02138 USA Tel: +1 617 395 4056 Fax: +1 617 354 6875 E-mail: [email protected]

©CAB International 2003. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Trees, crops and soil fertility: concepts and research methods edited by G. Schroth and F. L. Sinclair. p. cm. Includes bibliographical references (p. ). ISBN 0-85199-593-4 (alk. paper) 1. Soil fertility. 2. Agroforestry. 3. Soil fertility--Research. 4. Agroforestry--Research. I. Schroth, G. (Goetz) II. Sinclair, Fergus L. S596.7. T74 2002 631.4’22--dc21 2002006036 ISBN 0 85199 593 4 Typeset by Wyvern 21 Ltd, Bristol Printed and bound in the UK by Cromwell Press, Trowbridge

Contents

Contributors

ix

Foreword M. Swift

xi

1. Impacts of Trees on the Fertility of Agricultural Soils G. Schroth and F.L. Sinclair 1.1 Trees and the Development of Agriculture 1.2 Objectives and Structure of the Book 2. Economic Aspects of Soil Fertility Management and Agroforestry Practices A.-M.N. Izac 2.1 Introduction 2.2 Factors Influencing Farmers’ Decisions About Soil Fertility Management Practices 2.3 Hierarchy of Agricultural Systems as a Background to the Understanding of Farmers’ Constraints 2.4 Anatomy of a Decision at the Farm Scale and Economic Methods for Understanding such Decisions 2.5 Landscape and Global Scales: Soil Fertility and Agroforestry Trees as Part of Natural Capital 3. Designing Experiments and Analysing Data R. Coe, B. Huwe and G. Schroth 3.1 Synopsis

1 1 9 13

13 14 18 18 32

39 39 v

vi

Contents

3.2 3.3 3.4 3.5 3.6

Experimental Objectives, Treatments and Layout Fallow Experiments Measurements and Sampling Designs Analysing the Data Spatial Structure and Its Analysis

4. Soil Organic Matter G. Schroth, B. Vanlauwe and J. Lehmann 4.1 Synopsis 4.2 Methods for Total Soil Organic Carbon 4.3 Physical Fractionation Methods 4.4 Chemical Methods 4.5 Biological Methods

40 48 52 61 67 77 77 86 86 88 89

5. Soil Nutrient Availability and Acidity 93 G. Schroth, J. Lehmann and E. Barrios 5.1 Synopsis 93 5.2 Methods for Soil Nitrogen 104 5.3 Methods for Soil Phosphorus 112 5.4 Methods for Soil Sulphur 121 5.5 Methods for Potassium, Calcium and Magnesium in Soil 125 5.6 Methods for Soil Acidity 127 6. Decomposition and Nutrient Supply from Biomass G. Schroth 6.1 Synopsis 6.2 Methods for Biomass and Nutrient Input with Litter 6.3 Methods for Decomposition and Nutrient Release from Biomass 6.4 Measures of Resource Quality 7. Nutrient Leaching J. Lehmann and G. Schroth 7.1 Synopsis 7.2 Methods for Soil Solution Composition 7.3 Tracer Methods for Nutrient Leaching 7.4 Dyes as Tracers for Preferential Flow Paths 8. Nutrient Capture G. Schroth and J. Lehmann 8.1 Synopsis 8.2 Tracer Methods for Nutrient Uptake

131 131 140 143 148 151 151 158 163 165 167 167 174

Contents

9. Nutrient Exchange with the Atmosphere G. Schroth and J. Burkhardt 9.1 Synopsis 9.2 Methods for Atmospheric Nutrient Inputs 9.3 Methods for Nutrient Losses from Burning 10. Soil Structure M. Grimaldi, G. Schroth, W.G. Teixeira and B. Huwe 10.1 Synopsis 10.2 Methods for Soil Bulk Density 10.3 Methods for Aggregate Stability 10.4 Methods for Soil Porosity and Pore Size Distribution 10.5 Measuring the Role of Soil Organic Matter in Aggregate Stability 10.6 Soil Micromorphology and Image Analysis 11. Soil Water W.G. Teixeira, F.L. Sinclair, B. Huwe and G. Schroth 11.1 Synopsis 11.2 Methods for Soil Water Content 11.3 Methods for Soil Water Potential 11.4 Methods for Soil Hydraulic Properties 11.5 Estimating Topsoil and Subsoil Water Use with Stable Isotopes 12. Root Systems G. Schroth 12.1 Synopsis 12.2 Methods for Studying Root Distribution 12.3 Distinction of Roots of Different Species 12.4 Methods for Root Dynamics, Production and Turnover 13. Biological Nitrogen Fixation K.E. Giller 13.1 Synopsis 13.2 Microbiological Methods for Studying Rhizobia 13.3 Simple Methods for Determining Whether a Legume is Fixing Nitrogen 13.4 Isotope-based Methods for Measurement of Nitrogen Fixation 13.5 Estimating Nitrogen Fixation in Field Settings 13.6 Methods Based on Nitrogen Fixation Transport Products 13.7 Estimating Total Amounts of Nitrogen Fixation

vii

181 181 185 188 191 191 195 196 198 204 207 209 209 215 221 225 232

235 235 240 245 246 259 259 265 265 266 268 269 270

viii

Contents

14. Mycorrhizas D.L. Godbold and R. Sharrock 14.1 Synopsis 14.2 Inoculation Methods 14.3 Sampling of Plant Roots for Mycorrhizal Studies 14.4 Quantification of Mycorrhizas 14.5 Identification of Mycorrhizal Fungi 14.6 Estimation of Mineral Nutrient Uptake Through Mycorrhizas 15. Rhizosphere Processes D. Jones 15.1 Synopsis 15.2 Obtaining a Representative Sample of Rhizosphere Soil 15.3 Methods for Rhizosphere Soil Chemistry 15.4 Methods for Rhizosphere Biological Activity 15.5 Quantification of Root Carbon Loss into the Rhizosphere 15.6 Rhizosphere Mathematical Modelling 16. Soil Macrofauna P. Lavelle, B. Senapati and E. Barros 16.1 Synopsis 16.2 Sampling of Macrofauna: the TSBF Methodology 16.3 Other Sampling Methods 16.4 Manipulative Experiments

271 271 280 281 281 283 286

289 289 294 297 298 300 300 303 303 318 320 322

17. Soil Erosion M.A. McDonald, A. Lawrence and P.K. Shrestha 17.1 Synopsis 17.2 Quantitative Methods 17.3 Qualitative Methods

325 325 336 341

References

345

Index

423

Contributors

Edmundo Barrios, Centro Internacional de Agricultura Tropical (CIAT), AA 6713 Cali, Colombia Eleusa Barros, National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil Jürgen Burkhardt, Institute of Agricultural Chemistry, University of Bonn, Karlrobert-Kreiten-Str. 13, 53115 Bonn, Germany Richard Coe, World Agroforestry Centre, International Centre for Research in Agroforestry (ICRAF), PO Box 30677, Nairobi, Kenya Ken E. Giller, Plant Production Systems, Department of Plant Sciences, Wageningen University, PO Box 430, 6700 AK Wageningen, The Netherlands Douglas L. Godbold, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK Michel Grimaldi, Institut de Recherche pour le Développement (IRD), UR 137, Centre de Recherche d’Île de France, 32 ave Henri Varagnat, 93143 Bondy Cedex, France Bernd Huwe, Institute of Soil Science and Soil Geography, University of Bayreuth, 95440 Bayreuth, Germany Anne-Marie Izac, World Agroforestry Centre, International Centre for Research in Agroforestry (ICRAF), PO Box 30677, Nairobi, Kenya Davey Jones, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK Patrick Lavelle, Laboratoire d’Ecologie des Sols Tropicaux, 32 ave Henri Varagnat, 93143 Bondy Cedex, France Anna Lawrence, Environmental Change Institute, University of Oxford, 5 South Parks Road, Oxford OX1 3UB, UK ix

x

Contributors

Johannes Lehmann, College of Agriculture and Life Sciences, Department of Crop and Soil Sciences, Cornell University, 909 Bradfield Hall, Ithaca, NY 14853, USA Morag A. McDonald, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK Götz Schroth, Biological Dynamics of Forest Fragments Project, National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil Bikram Senapati, Ecology Section, School of Life Sciences, Sambalpur University, Jyoti Vihar, 768019, Orissa State, India Ruth Sharrock, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK Pratap Kumar Shrestha, Local Initiatives for Biodiversity, Research and Development (LI-BIRD), PO Box 324, Bastolathar, Mahendrapool, Pokhara, Nepal Fergus L. Sinclair, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK Wenceslau G. Teixeira, Empresa Brasileira de Pesquisa Agropecuaria-Amazônia Ocidental, CP 319, 69011–970 Manaus, AM, Brazil Bernard Vanlauwe, Tropical Soil Biology and Fertility Programme, PO Box 30592, Nairobi, Kenya

Foreword

Soil science benefits from the availability of a wide array of practicable methods. There are a number of very useful compendia in which sets of these methods are collected together to provide easy reference for the intending practitioner. The need for yet another handbook might therefore be questioned. This book, however, fulfils several needs that are not met in previous volumes. First and foremost it is unique in its format and purpose – being not a compendium of protocols but a reasoned discussion of the value and utility of different methods. Major advances in science are dependent on, and sometimes even driven by, the availability of suitable methods by which key questions or hypotheses may be answered. Progress may be said to be a product of the match between developments in concept and those in technique. The structure and content of this book are designed to review the ways in which current thinking in terms of the major problems of soil fertility can be methodologically attacked. As such the book should be useful not only in helping to solve problems but also in provoking new questions. The book is written within a particular context – soil fertility development under agroforestry. At first this may seem very specific and thus limited in appeal and application. But over the last decade or so agroforestry research has been one of the most influential in developing new insights into soil biology and fertility and thus provides a very suitable framework for review of progress. Furthermore, the influence of trees on soil is profound and of significance beyond agroforestry systems, so the book is likely to be of interest in the wider spheres of agriculture, forestry and ecological sciences. xi

xii

Foreword

The book covers a spectrum of soil methods broader than most soil science handbooks, combining the techniques of soil chemistry and physics with a wide scope of biological approaches and aspects of social science. This is reflective of the ways in which soil research has moved beyond its strict reductionist paradigm to embrace a more holistic and ecosystematic approach. The book comes at an appropriate time to review the progress of both concept and method in the slow march towards an integrated approach to land management. It should not only serve the purpose of directing enquiring researchers towards the useful approaches but also act as a stimulus for the next wave of soil fertility research. Mike Swift TSBF, Nairobi, Kenya

Editors’ Note This book was produced as part of a project of the International Union of Forest Research Organizations (IUFRO) on the development of manuals for research in agroforestry in association with the World Agroforestry Centre (ICRAF) and the Tropical Agricultural Research and Higher Education Centre (CATIE). Mike Swift, former Director of the Tropical Soil Biology and Fertility Programme (TSBF), has endorsed the book on behalf of the TSBF programme within the International Centre for Tropical Agriculture (CIAT). Götz Schroth received funding from the German Ministry of Education and Research through the German–Brazilian SHIFT programme and from the Brazilian National Council for Scientific and Technological Development (CNPq) while working on the book. The text was greatly improved by critical comments from several colleagues who read the whole manuscript or sections of it, especially Mike Swift, Ken E. Giller, Richard Coe, Michel Grimaldi, Bernd Huwe and Johannes Lehmann.

Chapter 1 Impacts of Trees on the Fertility of Agricultural Soils G. SCHROTH1

AND

F.L. SINCLAIR2

1Biological

Dynamics of Forest Fragments Project, National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil; 2School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK

1.1 Trees and the Development of Agriculture Trees are a natural component of most tropical landscapes, with the exception of very dry areas, tropical alpine ecosystems and a few other regions with extreme soil or climatic conditions which do not permit the establishment of woody perennial plants. In the humid tropics, trees are the dominant components of the natural vegetation, the tropical rain forests. With decreasing total rainfall and increasing length of the dry season, these are replaced by drier forest types and then by savannas. Although the density of trees decreases and their crown cover becomes more open with increasing aridity of the climate, trees are present and play an important role in natural ecosystems from the perhumid tropics almost to the fringe of the desert. Humans have had a pronounced influence on tropical ecosystems and their tree cover for a long time and continue to do so now. Fire and the axe, or its modern equivalent the chainsaw, are the main tools employed by farmers, planters and pastoralists to reduce tree cover and increase the availability of light and soil resources for their crops and pastures. Either tropical forests and savannas have been transformed by shifting cultivators into patchworks of swidden fields and different stages of fallow regrowth, or sedentary farmers have converted natural vegetation more permanently into crop fields, pastures, tree crop plantations or human settlements. Some of these areas may eventually be abandoned and revert to secondary forest or savanna vegetation, unless woody regrowth is prevented by recurrent fires, as in the Imperata grasslands of South-east Asia (Garrity et al., 1997). © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

1

2

G. Schroth and F.L. Sinclair

Although the rise in human populations has caused pronounced reductions in tree cover, trees remain an important element of most human-dominated landscapes throughout the tropics. Trees provide a wide range of important products and services that people in the tropics want and need. These range from firewood and construction materials, through many different fruits, nuts, medicines, gums, resins and fodder, to services such as shade, wind protection and aesthetic and spiritual value (Scherr, 1995). Tree cover also provides important habitat for both the conservation of wildlife and the utilization of many non-timber forest products that people harvest, including a number of valuable plants, fungi and game animals. Less visibly, but no less importantly, trees play a crucial role in maintaining and regenerating soil fertility through the action of their roots and litter. Tropical farmers are conscious of these different functions of trees and have protected, planted, selected and domesticated trees for thousands of years. Shifting cultivation systems in tropical forests depend on the regeneration of soil fertility under the forest fallow, which also provides game and a variety of products for collection. Shifting cultivators in Indonesia and other tropical regions have introduced tree crops such as rubber into their swiddens and have created secondary forests enriched with valuable trees (Gouyon et al., 1993). Farmers in West African savannas maintain valuable trees, which also resist periodical fires, in and around their fields, giving rise to a distinct, park-like landscape (Boffa, 1999). Planters of tree crops such as cocoa, coffee and tea maintain or establish shade trees to reduce pest and disease pressures and nutrient requirements of their crops and protect them from climatic extremes (Beer et al., 1998). Since many tree species can fulfil these functions, they may choose species that fix atmospheric nitrogen and produce large quantities of nutrient-rich litter and prunings, or valuable timber or fruit tree species, or any combination of these. Pastoralists value trees for the high nutritional value of the fodder from their leaves and fruits, which in seasonally dry pastures are still available when the grasses have dried out (Cajas-Giron and Sinclair, 2001). They may also prefer to plant trees as living fence posts rather than to replace timber posts every few years when they have been consumed by termites and fungi. Farmers in eastern and southern Africa have recently started to use short-term, planted fallows with legume trees to regenerate the fertility of their soils more rapidly than with natural fallows and to substitute for mineral nitrogen fertilizer, which is often too expensive for them to purchase (Kwesiga et al., 1999). These fallows may also produce valuable animal fodder. All these and many more practices that involve growing trees in some form of spatial or temporal combination with crops or pastures are known as agroforestry. They have drawn substantial interest from scientists and development agencies during recent decades, recognizing the fact that

Impacts of Trees on Fertility

3

trees can play an important role in income generation and food and fuel security for resource-poor rural households, while underpinning the sustainability of their farming systems (Cooper et al., 1996). This recent upsurge in interest in agroforestry may give the impression that it is a new science. This is not the case, as interactions between tree crops and shade trees, for example, have been studied by agronomists for more than 100 years and concepts of nutrient cycling, which are still relevant, were developed early on (see, for example, Lock (1888) on shade trees for coffee in Sri Lanka). Agroforestry is thus a relatively recent word for a much older science and a very old practice. However, efforts to make better use of trees in rural development had (and sometimes still have) to overcome initial antipathy between agriculture and forestry, institutionalized in government departments, research centres and educational establishments, which have led to an arbitrary separation of research and administration of forests and farming.

What is agroforestry? There are numerous definitions of agroforestry which stress different aspects of and expectations about the integration of trees in farming landscapes (see, for example, Huxley, 1999). Following the predominant definition over the past two decades, agroforestry is a set of land use practices that involve the deliberate combination of woody perennials including trees, shrubs, palms and bamboos, with agricultural crops and/or animals on the same land management unit in some form of spatial arrangement or temporal sequence such that there are significant ecological and economic interactions among the woody and nonwoody components. (Sinclair, 1999)

Traditionally, the focus of agroforestry research has been on interactions between trees and other components of a system, such as crops, soil and climatic factors, on the scale of an individual field or a small section of a landscape. This is gradually changing as better understanding of small-scale processes enables researchers to scale up their results, and as the functions of tree cover manifest at landscape, regional and global scales, as a result of larger-scale patterns and processes, become the focus of research interest. These functions include water and nutrient cycling on the catchment scale, carbon dynamics in the soil–plant–atmosphere system and biodiversity. Also, agroforestry practices do not exist in isolation, but interact with other land uses across landscapes. A farmer maintaining a forest garden or shaded tree crop plantation may also have swidden or irrigated rice fields and pasture which occur together within

Type of tree cover

Examples

4

Predominant land-use objective

Components

Trees on cropland – 2 Plantation tree crops – 2.1.1/2.2.3 Trees and crops silvoarable/agrosilviculture Crops on tree-land

Plantation forest/trees

Cropping phase of taungya – 2.1 Orchards/tree gardens – 2.1

Natural forest/tree cover

Forest gardens – 2.1.2 Cropping phase of shifting cultivation – 2.1.2

Trees on pasture/rangeland – 2 Plantation tree crops Trees and pasture silvopastoral

Pasture on tree-land

Plantation forest/trees

Forest/plantation grazing – 2.1.1 / 2.2.3

Natural forest/tree cover

Fodder banks – 2.2.5

Trees as fodder Trees and animals (not pasture)

Trees and poultry Trees and pigs Trees and fish Silk production

Trees and insects

Trees and tree crops

Multipurpose utilization of natural forest/tree cover

Primary or secondary forest

Improved fallow Hunting/collecting nontimber forest products

Trees on tree-crop land – 2

Fig. 1.1. Primary classification of agroforestry practices based on components, predominant land-use objective and the type of tree cover. Italics indicate examples of agroforestry practices, numbers refer to the place in Fig. 1.2 where classification continues (adapted from Sinclair, 1999).


Fallow phase of shifting cultivation Forests/trees


5

the same landscape and influence the characteristics of this landscape. This wider focus of agroforestry research is reflected in a scale-neutral definition of agroforestry as simply ‘where trees and agriculture interact’ (Sinclair, 1999). Within this wider view of agroforestry, the landscape scale is emerging as a critical unit of analysis (Sinclair, 2001). This is the scale at which ecological processes such as the presence and dispersal of fauna and flora, water and nutrient flows, microclimate, and pest and disease dynamics are significantly influenced by trees. In many fragmented landscapes trees on farms, including those shading tree crops or that occur as remnants in crop fields or pastures or in riparian corridors, provide key elements of the tree cover that determine landscape characteristics. Strategic placing of trees in the landscape may prevent, enhance or direct flows of soil, water, nutrients, fire and organisms across landscapes (van Noordwijk et al., 1999). Where there is a mosaic of agriculture and forest, interactions between these land uses determine such important environmental functions as the water yield of catchments and landscape-scale biodiversity 2.1.1 Regular

2.1 Dispersed trees in fields/on range

2.1.2 Irregular

2.2.1 On boundaries 2 2.2.2 On contours

2.2 Zoned

2.2.3 In rows

Live fences Boundary planting Windbreaks/timber belts On terraces On bunds As barrier hedges (contour hedgerows) Hedgerow intercropping/alley cropping Production hedges

2.2.4 Clumped Fodder banks 2.2.5 In blocks Farm woodlots

Fig. 1.2. Secondary classification of agroforestry practices based on density and arrangment of the tree component (adapted from Sinclair, 1999).

6


(Guindon, 1996; Bruijnzeel, 1997). In the future, efforts to understand and develop the role of trees on farms will increasingly focus on landscapes. In order to facilitate communication, the many ways in which farmers use trees within their farming systems have been classified into several major types of practice, on the basis of the components that are involved, the type of land on which they occur and the type of tree cover involved (Fig. 1.1). These major types of practice can be further classified in terms of the density and arrangement of the tree component (Fig. 1.2). This results in defining groups of practices which share important ecological and managerial characteristics. Lists and descriptions of common agroforestry practices can also be found in standard agroforestry texts (Nair, 1993; Young, 1997; Huxley, 1999).

Trees and soil fertility Whatever the reasons farmers have for planting or protecting trees in a specific case, they nearly always fulfil several functions simultaneously. Trees may have been planted on a hillslope to produce timber or fruits, but they may also protect the soil from being eroded. Trees planted or retained for fodder are often nitrogen-fixing and may improve nitrogen availability in the soil. Similarly, trees that have been allowed to regenerate in a riparian zone because of environmental regulation, for firewood production or simply an appreciation of their scenic value may also filter nutrients from runoff water, thereby retaining them in the land-use system and protecting the river from eutrophication. This book is about the effects of trees on soil fertility. Soil fertility is defined here as the ability of a soil to serve as a suitable substrate on which plants can grow and develop. Fertile soils facilitate root development, supply water, air and nutrients to plants, and do not have pest and disease burdens that result in catastrophic impacts on the plants that are being grown. Maintaining soil fertility is the basis of all forms of sustainable land use, that is, land use that remains productive in the long term. If fertility has fallen below a critical level through long-term agricultural use without replacement of nutrients or as a result of erosion, or if it is naturally very low, the replenishment of soil fertility may be a precondition for productive agriculture. Tropical soils are not generally infertile, but infertile soils are very common in the tropics. Moreover, recent research has provided evidence that soil fertility is decreasing in many farmed areas in the tropics. Nutrient budgets which were established at different spatial scales in sub-Saharan Africa, Ecuador and small farms in Costa Rica showed net nutrient losses from agricultural soils because of off-take in crop yields, leaching, erosion,


7

runoff and gaseous losses, which were not matched by nutrient inputs from mineral and organic fertilizers, atmospheric deposition, biological nitrogen fixation and, in flooded or irrigated areas, sedimentation (de Koning et al., 1997; Smaling et al., 1997; Stoorvogel and Smaling, 1998). Negative nutrient budgets like these are especially threatening for the fertility of soils whose nutrient stocks are already small, such as sandy soils with low organic matter contents, which are widespread in African savannas, or acid soils, which occupy vast areas of the humid tropics of Latin America, Africa and Asia (von Uexküll and Mutert, 1995). Accordingly, long-term fertility studies on farmers’ fields in African savannas have revealed evidence of widespread chemical and physical soil degradation, including negative soil organic matter and nutrient balances, although these have not always immediately been translated into declining crop yields (Pieri, 1989). Low and declining soil fertility are recognized by many tropical farmers as major constraints to agricultural production (Smaling et al., 1997). It can be expected that projected growth of human populations in tropical countries will further aggravate these problems, especially when population pressure increasingly obliges farmers to cultivate fragile and naturally infertile soils which are particularly prone to degradation. Both scientific research and farmers’ observations clearly point to the need to improve current farming practices with respect to their ability to increase and sustain soil fertility and agricultural productivity. The green revolution attempted to increase agricultural productivity in the tropics through increased inputs of mineral fertilizers, pesticides and new crop varieties. Although the successes were sometimes spectacular on relatively fertile land with good infrastructure, large numbers of small farmers in marginal environments were bypassed by these developments because they could not afford the necessary investment in new seeds and chemical inputs. Gradually, it has become understood that more accessible means are needed to enable many small farmers to feed their families and raise their living standards, and it has been suggested that this is most likely to come about from a thorough understanding of the biological bases of soil fertility (Woomer and Swift, 1994). Trees with their numerous beneficial effects on soil fertility play an important role in this strategy. The fundamental assumption in agroforestry, that the integration of trees into farming systems and landscapes can increase soil fertility, productivity and sustainability, was initially based mainly on the observation that soil under forest vegetation generally remains fertile and that tree fallows are able to regenerate degraded soils, as occurs in shifting cultivation systems (Nair, 1984). Subsequent scientific research has increasingly produced insights into the mechanisms through which trees improve soil fertility, though many key processes have still not been fully quantified. Agroforestry practices have been shown to influence chemical, physical and biological components of soil fertility. Trees can improve the

8


nutrient balance of a site both by reducing unproductive nutrient losses from erosion and leaching and by increasing nutrient inputs through nitrogen fixation; they can improve soil structure, water-holding capacity and crop rooting volume; and they can increase the biological activity in the soil by providing biomass and a suitable microclimate. However, a better understanding of the interactions between trees and soils has also helped to keep expectations at a realistic level and to recognize what agroforestry can and cannot achieve. Many commonly measured nutrient fluxes in agroforestry systems are part of the internal nutrient cycling within a system and do not change its overall nutrient budget. If a site is deficient in nitrogen and phosphorus, leguminous trees may be able to increase the availability of nitrogen through biological nitrogen fixation, but phosphorus may have to be added from external sources. It has also become increasingly clear that the intensity with which trees and tree–crop associations influence soil fertility differs widely between agroforestry practices, even if the processes are similar in principle. For example, the nitrogen fixation and biomass production of relatively few leguminous trees dispersed in a crop field will be much less than those of a closed stand of these trees in a planted fallow. Also, most trees will take up some of their nutrients from the subsoil and deposit them in surface soil through leaf litter and root decay and thus act as a nutrient pump. However, for some combinations of species managed in particular ways, this process may be important, whereas for others the effects may be too small to be of much consequence. When designing or improving agroforestry techniques, it is therefore important that the technique is matched with the fertility problems that are seen as priorities at a given site, rather than assuming that every type of agroforestry will improve soil fertility in general. Matching an agroforestry technique to the biophysical aspects of a site is necessary but not sufficient to ensure adoption; it also has to be compatible with the views, experiences, traditions and economic capacities of the farmers. Beneficial effects of trees on soil fertility are often only perceptible after several years and small farmers often cannot afford to invest in tree planting and tending without receiving an immediate return. Some techniques may require more time for pruning or biomass transport than the farmer can afford, especially when these activities are necessary at times when sowing, weeding or harvesting of crops are more urgent needs. It is also important to recognize that sustainable production from the same piece of land on the basis of stable soil fertility is not always a primary objective of the land user. Where the effort for clearing a new piece of land is smaller than that of maintaining the productivity of the already cleared plot, or where land clearing is a way of establishing ownership rights or is advantageous for other reasons, investments in sustainability may not have a high priority. Similarly, where runoff and


9

erosion from hillslopes benefit valuable crops in the valley below, soil degradation on the slopes may be seen as an acceptable price to pay for high productivity in the valley. It is thus clear that progress in agroforestry depends on a thorough understanding of both the biophysical and socioeconomic dimensions of farming systems at a range of scales.

1.2 Objectives and Structure of the Book This book provides an overview of the principal concepts that have been developed over the past decades concerning the effects of trees on soil fertility and an in-depth discussion of the methodological approaches that are appropriate to their study. It has been written mainly for researchers and students interested in tropical agroforestry. Case studies, examples and references have been taken mostly from the tropical agroforestry and ecology literature. However, although temperate agroforestry differs in its socioeconomic context from that in the tropics, the general biophysical processes are similar, and so most of the information presented here is also relevant to temperate conditions. Moreover, trees and soils in agroforestry are not fundamentally different from trees and soils in forests and especially in savannas, and we expect that students and researchers in forestry, agronomy and ecology will also find much useful information in the following chapters. Included in the book are economic, chemical, physical and biological aspects of soil fertility. Because of the integrated presentation of theoretical concepts and research methodologies, the book is particularly suitable for people who intend to do practical research on the interaction between trees and soils. It will be most useful to those who already have some basic knowledge of both agroforestry and soil science, although there are comprehensive references to the literature in these areas from which such an understanding could be gained. The chapters begin with synopses that are followed by methods sections. The synopses provide concise statements of essential background knowledge and outlines of the principal hypotheses and research results relevant to the topic at hand, and suggest key areas for future research. These sections are intended to help researchers in the identification of suitable, worthwhile topics for their research, and for students to see individual research results within the context of the larger body of knowledge and hypotheses relating to tropical soil fertility and agroforestry. The methods sections not only include those methods that are currently used in agroforestry, but also methods whose utility has been demonstrated in other fields and that could and should be applied in agroforestry studies in the future.

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This is not a methods book in the conventional sense, as it does not provide detailed field and laboratory descriptions of research methods. It has been produced to augment rather than to replace established methods texts, such as Tropical Soil Biology and Fertility: a Handbook of Methods (Anderson and Ingram, 1993), by furnishing a broader discussion of the scientific background of soils research in tropical agroforestry and the range of available research methods. The book also provides extensive reference to the relevant methodological literature, both from agroforestry and from soil science in general. The book starts with a chapter on the economics of soil fertility management and agroforestry practices, in which concepts for the analysis of farmers’ decisions are presented and consequences for the adoption or non-adoption of agroforestry practices are discussed. Attention is drawn to the substantial public benefits at national and global scales that derive from the ecosystem services provided by trees. At present farmers often bear the costs of these public goods, which results in a level of agroforestry adoption that may be lower than is desirable from a societal point of view (Chapter 2). Chapter 3 presents some general grounding in appropriate methods for experimentation, sampling and data analysis for soil fertility research in agroforestry. A section on fallow experimentation is included because of the particular requirements of working with rotational systems that have distinct temporal phases and their present importance in agroforestry research. There is also a section on geostatistics, a tool that has not yet been widely used in agroforestry research but may become increasingly important in the future because of its usefulness in the analysis of spatial heterogeneity. Chapter 4 discusses the dynamics of organic matter in tropical soils as influenced by agroforestry and other land-use practices. The following four chapters are dedicated to different aspects of nutrient cycling. The synopsis section of Chapter 5 discusses the principal hypotheses concerning how trees may affect nutrient cycles and introduces the concepts of competition, complementarity and facilitation. This discussion provides a framework within which the macronutrients nitrogen, phosphorus, potassium, calcium and magnesium as well as soil acidity are discussed. The next three chapters treat nutrient cycling processes of particular relevance to agroforestry: decomposition and nutrient release from biomass (Chapter 6), nutrient leaching (Chapter 7) and nutrient capture (Chapter 8). Atmospheric nutrient inputs and nutrient losses into the atmosphere through fire are the topics of Chapter 9. Physical soil fertility is treated in the following two chapters on soil structure (Chapter 10) and soil water (Chapter 11). Five chapters are dedicated to biological aspects of soil fertility: root systems (Chapter 12), biological nitrogen fixation (Chapter 13), mycorrhizas (Chapter 14), rhizosphere processes (Chapter 15) and soil fauna (Chapter 16). Soil erosion is the topic of the last chapter (Chapter 17). Throughout the text


11

some effort has been made to demonstrate that the chemical, physical and biological components of soil fertility strongly interact. Topics related to soil fertility and agroforestry that have not been included in this book include salinization and waterlogging, which have been addressed in several contributions to a recent symposium (Lefroy et al., 1999); soil-related pest and disease problems, which are still a neglected field in agroforestry despite recent progress (Desaeger and Rao, 1999; Duponnois et al., 1999); and allelopathy, on which a large literature exists (e.g. May and Ash, 1990; Ramamoorthy and Paliwal, 1993; Conger, 1999), although there remains a paucity of information about its practical importance in agroforestry.

Chapter 2 Economic Aspects of Soil Fertility Management and Agroforestry Practices (A.-M.N. IZAC) World Agroforestry Centre, International Centre for Research in Agroforestry (ICRAF), PO Box 30677, Nairobi, Kenya

2.1 Introduction Analyses of adoption of soil fertility management and agroforestry practices by farmers in tropical countries are relatively sparse. They provide a patchwork picture of cases where adoption occurred relatively rapidly and was quite widespread and other cases where adoption did not occur on any significant scale. This chapter deals with the economic concepts that can be used by scientists interested in increasing the probability of adoption of soil fertility management practices by farmers. It is stated in Chapter 1 of this book that: ‘When designing or improving agroforestry techniques, it is ... important that the technique is matched with the fertility problems that are seen as priorities at a given site, rather than assuming that every type of agroforestry will improve soil fertility in general.’ The goal in this chapter is to present economic concepts for matching soil fertility interventions to both farmers’ constraints and objectives and those of society, in order to increase their likelihood of adoption. The chapter highlights key economic concepts and processes that biophysical scientists should find helpful in putting their own work into the broader context of the farmer’s economic environment. It does not attempt to provide detailed discussions of economic approaches to the assessment of agroforestry practices for professional economists (economists are not the major intended audience for this book). Furthermore, such detailed discussions are not readily available in the literature, and therefore may constitute a gap which needs to be filled. Declining soil fertility is acknowledged as a problem by the vast © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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majority of farmers who experience it on their farms (see Chapter 1). It is, at the same time, a problem for society as a whole as it is related to issues of agricultural sustainability, soil biodiversity, carbon sequestration and watershed functions. This chapter focuses on the decision-making processes that determine whether an agroforestry intervention will be adopted by farmers and whether ensuing levels of adoption will be socially optimal. These processes are analysed within the framework of ecological economics, which differs substantially from mainstream economics. The emerging field of ecological economics addresses the relations between ecosystems and social and economic systems. Social and economic systems are viewed as subsystems of the biosphere and thus as wholly dependent upon ecological–economic interactions. This is in direct contrast with the conventional or mainstream economic approach, which considers that all phenomena are subsumed within and obey the rules of an economic system. In what follows, factors affecting farmers’ decision-making processes regarding adoption of soil fertility and agroforestry practices are first highlighted. A brief discussion of agricultural systems hierarchy serves to set the context within which farm-scale decision-making processes are analysed. Finally, economic processes relevant at the regional and global scales are discussed.

2.2 Factors Influencing Farmers’ Decisions About Soil Fertility Management Practices The decision to adopt a given soil management technology is made by an individual farmer on the basis of a number of factors which the farmer integrates into a framework driven by the farmer’s production objectives. Social scientists who have analysed farmers’ decision making in the tropics have shown that farmers think in a systemic fashion. Decisions regarding a given field or a given practice, such as soil fertility management and agroforestry, are thus not made in isolation. These decisions are made within the context of the whole farm and of the totality of the resources and assets available to the farmer. These resources and assets include: (i) labour (family labour plus hired labour if sufficient cash is available); (ii) cash to buy fertilizer and other chemicals; (iii) their entire landholding and the different fields comprising it; (iv) purchased assets such as implements, machinery, animal traction; (v) access to water (either on farm or off farm); and (vi) access to other off-farm resources (such as communal resources, forested lands and woodlots). Farmers focus on the trade-offs between the efforts they have to make to meet their production objectives (being able to produce enough food

Economic Aspects of Soil Fertility Management

15

for the family, being able to produce a surplus and to sell it) and the payoffs they expect from these efforts. They consider the totality of their holding and its various current and potential uses vis-à-vis their production objectives and weigh these in terms of the outcomes they expect when combining their resources into different practices (including soil fertility management). Farmers also have to factor into such decisions different groups of markets and different types of customs. They need to consider markets for agricultural commodities (local, regional, national and international if relevant), since the sale of their surplus production, over and above what they need to produce to feed their family, depends on these markets. They also have to take into account markets for inputs. These determine the costs they will have to incur for their use of: (i) labour, be it hired or family labour; (ii) land, especially if they rent some of their fields; (iii) capital, especially if they borrow money; (iv) implements and machinery; and (v) water and other resources. The level of income of farmers is determined by these two groups of markets. Finally, the actual purchasing power of this income is dictated by markets for consumer goods (such as clothing and medicines) and by government policies regarding public infrastructure (such as means of transport and their costs, costs of education, health and electricity). Social customs and norms influence a number of the elements that farmers need to integrate in their decisions. These customs and norms determine farmers’ access to many natural resources, as well as human labour, through the prevailing land and tree tenure system, and the regulations concerning access to off-farm resources (water, communal resources such as forests and woodlots), and access to non-family labour. In some societies in western Africa for instance, rules of access to labour are extremely complex and based on what anthropologists call reciprocal relations. It is important for scientists undertaking research on the development of soil fertility management practices to be cognizant of the fact that farmers’ decisions to adopt such practices are complex and driven by the conjunction of the above factors. Scientists need to understand the basic principles at play in such decisions in order to design interventions that match farmers’ constraints and are therefore adoptable. The various types of factors influencing farmers’ decisions are depicted in Fig. 2.1. Figure 2.1 also illustrates the fact that these factors are determined at different spatial scales. Although some of them are clearly determined at the farm scale (e.g. family labour and soil fertility status of the fields) others are determined at the landscape and community scale (e.g. access to off-farm resources) or at the regional and national scale (e.g. government policies concerning rural infrastructure and markets for consumer goods).

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.. .. ..

.. .. ... .

Social–political parameters political system social customs land and resource tenure demography infrastructure (transport, credit, education, health)

Economic parameters markets for inputs and outputs degree of integration in markets non-farm income technologies available existence of a surplus from previous year endowment in family labour, land, capital

.. .. ..

Ecological parameters climate soils hydrology pests weeds topography

Farmers’ production objectives

.. ..

Farmers’ decision making what to produce when to produce it how to produce it (including soil fertility management practices)

Outputs produced quantity and quality (crops, animals, trees…)

Environmental impacts depending upon practices chosen: soil erosion, acidification, water pollution, deforestation, decreased genetic diversity, or : enhanced natural resource base, sustainable production

Fig. 2.1. Context of farmers’ decisions to adopt.

Finally, Fig. 2.1 shows that a decision to use specific practices, as per these factors and per the farmer’s production objectives, have specific outcomes for the farming household and may also have off-farm consequences. The off-farm consequences include, for instance, the forest degradation that would result from farmers deciding to ameliorate the effects of decreasing soil fertility on their fields by transferring forest soil to their land, as occurs in parts of India. Another example would be the increased water sedimentation for downstream farmers resulting from a decision to clear secondary fallow land for agriculture by upstream farmers in northern Thailand. The purpose of this section is to debunk some often-held myths about farmers’ decisions concerning soil management in tropical countries. The following points are stressed. First, farmers in tropical countries are as rational in their decision making as any other persons or stakeholders in any country. Secondly, they generally have a different perspective from


17

that of scientists when considering improved soil management practices. Theirs is a system perspective whereas scientists often adopt a reductionist perspective when designing technologies. This may explain why rates of adoption of these technologies by farmers are often a disappointment to scientists who have not factored into their technology design all the relevant constraints faced by the farmers. Thirdly, farmers’ decisions to

SUPRAREGIONAL SYSTEM Large geographical and economic units

Climate, geology, etc.

National economic policies and institutions

International trade

REGIONAL SYSTEM Natural resources endowments, human resources endowments

Industrial and urban regions

Rural regions

Regional institutions, policies

VILLAGE/LANDSCAPE SYSTEM Topography, pest population, etc.

Farming households

Non-farming households

Social norms and customs, roads, infrastructure

FARMING SYSTEM Various inputs

Decisionmaking units

Livestock production

Agroforestry production

Crop production

AGROFORESTRY SYSTEM Weeds

Insects

Pathogens

Crops

Soil

Water

SOIL MANAGEMENT SYSTEM Mulching

Tillage

Manuring

Management

Fig. 2.2. Hierarchy of systems within which farmers make decisions.

Trees

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adopt are constrained by various factors, determined at different spatial scales; likewise these decisions to adopt can have consequences at spatial scales beyond the farm scale.

2.3 Hierarchy of Agricultural Systems as a Background to the Understanding of Farmers’ Constraints Figure 2.2 represents a spatial hierarchy of agricultural systems within which farmers’ decisions are made. The highest level in the hierarchy, supraregional systems, occupies the largest land area and can transcend national boundaries. Macroeconomic processes, as well as certain geological processes, are best understood at this level. The lowest level (soil systems) covers the smallest spatial unit and is the level at which specific biological processes such as nutrient uptake may be investigated. A basic rule in systems theory is that systems at level n are constrained and controlled by systems at level n + 1, and in turn they constrain systems at level n – 1 (Allen and Starr, 1982). Farmers (operating at the farming system level) thus have to take the environment provided by the village or landscape level as a constraint in their decisions regarding soil fertility practices. Likewise, the village or landscape scale is constrained by regional system variables that operate themselves within the confines of supraregional variables. Consequently, farmers integrate a wide range of ecological, social and economic parameters belonging to levels higher than the farming system in their decisions to adopt soil fertility management and agroforestry practices. Decisions made at the farming system scale have repercussions at the same scale, as well as at lower and higher scales in the hierarchy. These are mediated through various economic and biological processes, such as nutrient cycling and the market mechanism. Because these processes transcend farm boundaries, it is helpful to establish a distinction between the economic processes that occur at the farm scale and those that are manifest at the landscape or watershed and global scales. Even though soil fertility management and agroforestry practices are very localized interventions on farmers’ fields, it is essential for scientists to realize that the key processes of relevance to their adoption occur at the farm, regional and global scales.

2.4 Anatomy of a Decision at the Farm Scale and Economic Methods for Understanding such Decisions The decision by an individual farmer to adopt a given soil fertility management or agroforestry practice is made within the frame of the


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factors shown in Fig. 2.1. More specifically, such decisions are based on farmers’ production objectives and farmers’ perceptions of the advantages and disadvantages of a given practice. These advantages and disadvantages are all the monetary and non-monetary costs and benefits of a practice, as perceived by farmers. All these costs and benefits are relative to, or determined by, the existing land tenure system and markets. For instance, Unruh (2001) has shown that in post-war Mozambique, where land rights are unclear and ambiguous, land disputes are very common and costly. Agroforestry trees, in particular older cashew trees (Anacardium occidentale), are, however, considered as evidence of land ownership. In such an institutional context, these trees have a substantial non-monetary benefit; they serve to clearly establish stronger land claims for farmers (Unruh, 2001). Notions of costs and benefits are thus highly relative concepts in economics. What is a benefit in a given institutional context, such as trees that establish land rights in Mozambique, may not be so in countries where land rights are unambiguous. In what follows, the basic economic principles and processes at play in farmers’ decisions to adopt or not to adopt are analysed, and examples from actual economic analyses are given to illustrate the argument. The ‘real-world’ costs and benefits of a given agroforestry practice in a given location are highly site specific and variable across farms. This is a direct consequence of the complex range of factors that determine the value of each cost and benefit. It is thus very important for scientists interested in understanding farmers’ decisions to be aware of the economic processes that guide these decisions. Without such an understanding, empirical evidence of the profitability (or lack thereof ) of specific agroforestry options may appear very confusing.

Principles of private cost–benefit analysis and net present value One of the simplest ways to assess these costs and benefits is to use private cost–benefit analysis. This analysis consists of comparing the flows of benefits and costs generated over time by the adoption of a given agroforestry practice. Only the costs and benefits that are relevant to a given farmer are taken into consideration, and they are valued at the market prices faced by the farmer. No attempt is made to adjust costs and benefits for market and government failures. Nor is any attempt made to take social costs and benefits (externalities and environmental benefits) into consideration. Pagiola (1994) and Sugden and Williams (1978) provide excellent introductions to cost–benefit analysis for non-economists. Gittinger (1982) is one of the classic texts on cost–benefit analysis, along with Mishan (1976). Both provide much detail about the approach, its basic

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assumptions, and various methods of computing costs and benefits. Both texts are written for professional economists. The basic principle in private cost–benefit analysis is simple. The net value, in today’s dollars, of the future flows of benefits and costs of an agroforestry option is computed. In economic language, this is called the net present value of an agroforestry option. This net present value (NPV) is defined as: NPV =

n

Â t =1

Bt – Ct

(1 + r)

t

(2.1)

where: Bt = benefits in year t; Ct = costs in year t; r = discount rate, that is, the rate at which benefits and costs incurred in year t have to be discounted to arrive at their value in today’s dollars; and n = number of years during which the agroforestry option will generate benefits and costs. In what follows, the concepts of costs and benefits are further discussed, as well as the concept of discount rate.

Monetary and opportunity costs of adoption The generic soil fertility and agroforestry practices that are discussed in the other chapters of this book include improved fallows (tree–crop rotations), tree–crop associations (such as hedgerow intercropping and shaded perennial crops), contour hedgerows and boundary plantings. Each of these options entails various monetary and non-monetary costs for farmers. Therefore: Ct = mct + oct

(2.2)

where mct = sum of monetary costs in year t, and oct = sum of opportunity costs in year t. The monetary or out-of-pocket costs (mct) are quite straightforward. They consist of: (i) the costs of buying seeds and/or seedlings (or the costs of having an on-farm nursery for the tree seedlings), and whatever other inputs such as fertilizer are used, and (ii) the costs of hiring additional labour (if family labour is insufficient) for planting the trees, weeding the trees, pruning them, and applying or incorporating tree prunings to the soil (see Table 2.1). The most meaningful way of assessing these costs is on the basis of on-farm experiments or under actual farming conditions, through on-farm monitoring of farmers’ practices. Monitoring the use of hired labour throughout the year, and throughout the lifespan of an agroforestry option is not an easy task, and is costly when this lifespan is long. There are therefore relatively few examples of private cost–benefit


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Table 2.1. Generic on-farm costs of agroforestry practices. • Monetary costs of additional hired labour for planting tree seedlings/seeds, pruning the trees, weeding the trees (most agroforestry options) • Monetary cost of purchase and transport of seeds and seedlings and fertilizer (if applied; most agroforestry options) • Opportunity cost of not using a field or part of a field in another way (for improved fallows, intercropping, alley cropping and trees on contour ridges) • Opportunity cost of family labour used (forgone participation in other farming activities; most agroforestry options) • Opportunity cost of possible losses in germination of crops (for agroforestry options requiring the application of tree residues to crops) • Opportunity cost of lower crop yields due to competition from trees (for intercropping, alley cropping and parklands)

analyses that use such ‘real-world’ data. Many analyses rely on on-station experiments and on extrapolations of on-station data to on-farm conditions. Dewees (1995) provides an interesting analysis of the costs to farmers in Malawi of intercropping maize with Faidherbia albida, and of alley cropping Leucaena leucocephala with maize. The non-monetary or opportunity costs (oct in Eq. 2.2) are not actual disbursements for farmers, but are nevertheless very real costs which farmers take into consideration in their decisions. They consist of all the opportunities for generating benefits that a farmer gives up when choosing a given agroforestry option (see Table 2.1). The appropriate baseline for such comparison is the best alternative practice that the farmer could have chosen in lieu of this agroforestry option. If there is an alternative practice that would bring higher benefits to the farmer than agroforestry, then it is highly unlikely that the farmer will choose agroforestry, unless agroforestry also brings about significant intangible social benefits (such as increased social status in the community) for the farmer. Two principal types of opportunity costs of agroforestry options are those of labour and land. When family labour is used for planting hedgerows, for example, the opportunity cost is equal to the benefit that this family labour could have generated if instead of planting seedlings it had been engaged in the best alternative, such as planting or weeding food or cash crops. The opportunity cost of family labour can sometimes be a major constraint to the adoption of agroforestry practices because: •

•

the extra labour required for implementing some of these practices may be needed at a time of year when all household members are occupied in tasks given a high priority (e.g. production of food crops); and/or available family labour is decreasing in some areas due to HIV/AIDS; and/or

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in some countries such as Kenya there is a fairly strict division of onfarm responsibilities between females and males, so that even if female labour is plentiful some tasks such as pruning trees in hedgerows may not be done because male labour is insufficient (see Swinkels and Franzel, 1997, for details).

In such cases, private cost–benefit analysis in itself may not be sufficient in capturing these non-quantifiable constraints, and farmer surveys will be needed to complement the picture painted by private cost– benefit analysis. It could indeed be the case that a given agroforestry option, although economically profitable, is at the same time not really feasible, and therefore will have a low adoption rate, for the reasons just mentioned. Whereas monetary costs of adoption are simply assessed by their market value (e.g. cost of one seedling times number of seedlings purchased; hourly wages paid to hired labour times number of hours worked), opportunity costs are more difficult to assess. In the case of family labour, economic theory indicates that the prevailing wage rate times the number of hours worked by family labour should be used to assess this cost. This is because it is assumed that, if family labour had not been working on the farm, it could have been engaged working as hired labour on other farms. When rural unemployment is high, however, it is doubtful that it would have been feasible for family members to be thus employed in the agricultural sector. As a rule of thumb, the higher the rate of unemployment in an area, and the more difficult for family members to travel within this area (because of geographical isolation or bad transport networks), the lower is the wage rate that should be used to assess the opportunity cost of family labour (see Pagiola, 1994, for a clear discussion of these issues). The opportunity cost of an agroforestry option in terms of the land area it occupies (entire field or portion of fields) is easier to assess. It is equal to the benefits the farmer would have gained from the best alternative land use. This is generally the dominant or traditional land use in an area, such as the cultivation of maize with a very low level of fertilizer in the case of improved fallows in western Kenya. In such an example, the opportunity cost of the land under improved fallows is equal to the net benefits the farming household would have received from the traditional maize cultivation system on a per hectare basis, times the land area under fallow. In some instances, the best alternative to the agroforestry option under analysis may be a different soil fertility enhancement method that farmers could choose to use (e.g. green manuring with legumes). In this event, net benefits from manuring with legumes on a per hectare basis times the land area under fallow will be equal to the opportunity cost of a fallow practice.


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Not only are these various costs (represented in Table 2.1 and equal to Ct in Eq. 2.2) entirely borne by the farming household, but also the household must start paying for them from the moment a practice is chosen for implementation. In addition, some of these costs will recur over a number of years. In some of the few assessments of actual costs of adoption incurred by farmers published in the literature, total costs of adoption decrease very substantially between year one and the following years, and are, during the first years, significantly higher than the costs of implementing farmers’ current practices. This was the case, for instance, for hedgerow intercropping (maize with Leucaena leucocephala or Calliandra calothyrsus) in the highlands of western Kenya. The total costs of adoption to farmers (Ct) amounted, on average, to US$422 ha–1 during the first year and decreased, on average, to US$276 ha–1 during the second year (Swinkels and Franzel, 1997). This compared with total costs of US$296 for the traditional maize system during the first year and costs of US$271 during the second year. In another example, of improved fallows in Costa Rica (with Vochysia ferruginea and Hyeronima alchorneoides), actual costs (for clearing, planting, replanting, weeding, pruning, and seedlings and their transport) were around US$1330 ha–1 for the first year. They decreased to US$162 ha–1 during the second year and US$97 ha–1 during the fifth year of adoption (Montagnini and Mendelsohn, 1997).

Monetary and non-monetary (environmental and social) benefits of adoption The benefits of agroforestry practices, as they are perceived by the farmer, are generally more difficult to assess than their costs. And, furthermore, some of these benefits often start occurring after a while, so that there is a time lag between the moment when a household incurs costs in order to adopt an agroforestry practice and the moment when it starts receiving benefits from this practice. In parallel with costs, the benefits of adoption of a practice can be monetary and non-monetary (see Table 2.2). Therefore: Bt = mbt + ibt

(2.3)

where mbt = monetary benefits in year t; and ibt = non-monetary benefits in year t. There are two kinds of monetary benefits (mbt). The first is the value of the increased crop yields associated with a given agroforestry practice. In a maize-based improved fallow system for instance, this will be the market value of the increased maize yields following the improved fallow period. Such direct monetary benefits will occur over a period of time, that

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Table 2.2. Generic on-farm benefits of agroforestry practices. Timea Benefits

Year 1

Years 2–5

Monetary Increased crop yields + ++ Value of various tree products 0 + Non-monetary Increased resilience and sustainability of system through: Decreased risks of yield fluctuations with usual climatic variability 0 + Enhanced soil resource base 0 0 Enhanced capacity of system to adjust to exogenous changes without generating increased flow of pollutants 0 0 Increased biodiversity of soil biota 0 + Increased biodiversity of fauna and flora 0 0

Years 6–10

++ ++

++ +

+ ++ +

a The time scales shown are indicative. 0, no measurable benefit; +, ++, benefit is measurable and its intensity varies from low (+) to high (++).

is, until all residual effects of the improved fallow have been exhausted. To continue with our example, it is thus necessary to assess maize yields over time in the improved fallow system (as well as maize yields over time in the traditional maize system, since their market value will constitute the opportunity cost of the land under fallow). Nominal farm-gate prices of maize over the relevant period of time also need to be obtained or predicted. Even if the maize is grown for home consumption rather than for sale, the value of this monetary benefit of adoption will be evaluated as: tonnes of maize produced multiplied by the market (farm-gate) value of a tonne of maize, divided by the relevant number of years. The second type of monetary benefit is the market value of the diverse products that may be generated by an agroforestry practice. Examples of these products include indigenous fruits, timber, nuts, leaves that are used in food preparation, leaves, bark and seeds that have medicinal properties, poles, fodder, fuelwood. Proceeds from the sale of these products will contribute to household income and are thus a monetary benefit. If these products are not sold but are consumed by the household, their market value equivalent still represents a monetary benefit, since the household does not need to spend some of its income on the purchase of these products. In a very real way, these products directly contribute to the welfare of the household. Leakey and Tomich (1999) provide various examples of the market values of tree products from agroforestry systems,


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although these systems are not restricted to those that enhance soil fertility. For all such monetary benefits, the effects of an agroforestry option on the yields of various products, from crops to medicinal products, need to be quantified on a yearly basis over the relevant time period. The value of these yields is then obtained by assessing or predicting the farm-gate prices of these products. To use again the example of hedgerow intercropping in western Kenya, total monetary benefits increased from US$709 ha–1 during the first year to US$766 ha–1 during the second year (Swinkels and Franzel, 1997). This compared with total monetary benefits of US$707 ha–1 for traditional maize; these benefits remained at this level over time. The non-monetary benefits of agroforestry practices at the farm scale (ibt in Eq. 2.3) are particularly difficult to assess. They consist of all the improvements in soil and other ecological processes that are brought about by the agroforestry practices and are not captured by increased crop yields. In other words, to understand and assess these non-monetary benefits, one needs to first consider all the functions which enhanced soil fertility (such as increased nutrient stocks, enhanced efficiency of nutrient cycling) will have on a farm and which are over and above those already translated into increased crop yields. One such function that farmers, including resource-poor households, particularly value is the increased resilience and sustainability, and therefore the increased risk-buffering capacity, of their systems. Recent poverty surveys undertaken by the World Bank (Kanbur and Squire, 2000) show that the poorer a farming household, the more important it is to this household to have effective strategies for coping with risk (environmental, climatic, economic). Ability to manage risks and adapt to change is actually something which poor farmers rank as a priority concern, on a par with increasing their income (World Bank, 1994; Kanbur and Squire, 2000). The next section shows that one method of evaluating this increased risk buffering capacity of agroforestry options is to use a lower rate of discount (r in Eq. 2.1). There are many other examples of non-monetized benefits of agroforestry practices, such as increased soil biodiversity, decreased erosion, increased water infiltration and recharge of underground water table, improved soil structure, enhanced capacity of systems to adjust to change without generating increased flows of pollutants, enhanced carbon sequestration. Although some of these will eventually result in increased yields (e.g. improved soil structure), others will not have any measurable effects on crop yields (e.g. improved recharge of the underground water table). Biological scientists have not reached a consensus about how to assess these various functions of enhanced soil fertility. It would thus be unrealistic to assume that farmers will be aware of all these functions.

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Furthermore, even if they were aware of them, it is unlikely that they would factor all of them into their decisions to adopt. This is because some of these functions, such as increased soil biodiversity, are likely to be of no special interest to them. Finally, non-monetary on-farm benefits of adoption can include aesthetic value, habitat for welcome wildlife and shade. Farmer surveys have shown that such benefits can be valued very highly by farmers (Scherr, 1995). Table 2.2 presents generic categories of benefits of agroforestry systems for soil fertility improvements (Bt in Eq. 2.3) and illustrates a significant difference between the costs and benefits of adoption of such practices. This difference has been observed in a number of assessments of the costs and benefits of agroforestry practices (Current et al., 1995; Dewees, 1995; Montagnini and Mendelsohn, 1997; Swinkels and Franzel, 1997). Most benefits are cumulative over time and reach greater amplitude 3 or 5 years following adoption whereas costs have to be borne up front, as explained previously.

Farmers’ time horizons, discounting and balancing of the costs and benefits of adoption It was seen above that there are some fundamental differences between the costs and the benefits of agroforestry systems for soil fertility management at the farm scale. The costs are entirely borne by farmers and occur from the inception of adoption of a practice. The benefits are generally cumulative over time and low during the first years following adoption. In addition, farmers are unlikely to value some of the nonmonetized benefits such as increased soil biodiversity. This raises the issue of the time frame of relevance to farmers. Recall that the NPV of an agroforestry option is represented in Eq. 2.1. This equation reflects the fact that benefits (and costs) occurring at some future time do not have the same value today as they will have in the future. Indeed, if we are asked whether we prefer to receive US$100 today or in 3 years’ time, most of us will answer that we prefer to receive this sum today. This is because we can either spend the money now and immediately receive some enjoyment from it, or invest it to receive a greater amount in the future (US$100 plus interest over 3 years). The economic concept of a rate of discount (r in Eq. 2.1) is the measure which enables economists to translate future benefits and costs into today’s monetary worth. It is the rate at which we need to be compensated in the future for willingly accepting not enjoying a benefit today. It is also called the rate of time preferences (see Mishan, 1976, for a detailed discussion).


27

As can be appreciated by simply looking at Eq. 2.1, the choice of a specific value for r is extremely important. Indeed, the value of NPV is highly sensitive to the value of r. When costs are incurred in the present and medium term, whereas benefits occur over the medium and long term, the lower the rate of discount used in the analysis, the higher the resulting NPV, and vice versa. Eliciting farmers’ true rate of time preferences is a difficult task. Economic, psychological and social factors influence this rate. To arrive at a realistic rate through very carefully designed and in-depth interviews is a research project in and of itself. Consequently, in practice analysts use a rate, chosen on somewhat arbitrary grounds, that they assume reflects as well as possible the perspective of farmers in a given community. Resource-poor small-scale farmers in the tropics are highly vulnerable to risks (Kanbur and Squire, 2000) and generally concerned about their survival in farming from one year to the next. It is therefore likely that, even if they were aware of the medium- and long-term benefits of adoption, most farmers would value these future benefits relatively less than immediate benefits. This is because they occur over a period of time of little relevance to their immediate needs. In other words, the discount rate of relevance to farmers in tropical countries to be used in Eq. 2.1 is likely to be quite high (rates of around 20% are used in most studies). One empirical study of farmers’ discount rates suggests that Costa Rican farmers discount the future at a rate of 20–25% (Cuesta et al., 1994). Another study of smallholders in the Philippines indicates that 40% would be an appropriate rate of discount for these farmers (Nelson et al., 1998). It should be noted that, when an agroforestry option is seen by farmers as a method for managing risk (such as farmers in Rajasthan who cope with repeated droughts by the complementary use of woody perennials and annuals), then their implicit rate of discount decreases substantially (Arnold, 1997), and rates of 5–10% can be used. Given the difficulties associated with the choice of an exact rate of discount and the sensitivity of the resulting NPVs to this choice, a good analytical practice is to use a range of rates. The corresponding range of NPVs will highlight the threshold rate of discount at which an agroforestry option becomes privately profitable for farmers. The relatively few empirical assessments of the NPV of different agroforestry practices generally indicate a positive NPV. For instance, in the 21 agroforestry projects implemented in eight countries of Central America and the Caribbean that were reviewed by Current et al. (1995), NPVs of trees with crops, alley cropping, contour planting, perennial crops with trees, homegardens and taungya systems were all positive, whereas the NPV of woodlots was negative at a 20% discount rate, indicating net economic losses for farmers. In another study on the loess plateau in China, the NPV of an agroforestry intervention consisting of apple

28

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orchards on bench terraces was positive, and was also about 100% higher than the NPV of soil conservation practices without a tree component (Lu and Stocking, 2000). Comparing the NPV of an agroforestry option to that of alternative options for farmers is an important step in understanding decision making at the farm scale. Various studies suggest that agroforestry systems typically have higher NPV than current or traditional practices (e.g. Current et al., 1995, for Latin America and the Caribbean and the above example from China), with some notable exceptions in Africa (e.g. Dewees, 1995, for intercropping and alley cropping in Malawi; Drechsel et al., 1996, for improved fallows in Rwanda; Swinkels and Franzel, 1997, for alley cropping in Kenya). In addition to ensuring that the over time private profitability of an agroforestry option is compared to other options of relevance to farmers interests, it is also important to assess the sensitivity of this measure of profitability to changes in some variables. Sensitivity analysis consists of varying different parameters in Eq. 2.1, such as the price of labour, the yields and prices of the crops produced, to assess the effects of these changes on NPV (see Gittinger, 1982, for further details). The data used to assess these parameters have a built-in uncertainty, given the paucity of relevant secondary databases in tropical countries, the unreliability of existing data and the fact that primary data are costly and often difficult to collect (see Lele, 1991, for a discussion of these issues). It is therefore important to investigate the effects of changing the values of these parameters, within reasonable bounds, on the results of the analysis to better appreciate the robustness of the profitability of an agroforestry option. The study by Current et al. (1995) indicates that the positive and high NPVs obtained for the vast majority of agroforestry practices in various Latin American and Caribbean countries were most sensitive to changes in yields and product prices. In the African context, it would appear that results of NPV are most sensitive to changes in adoption costs and yields (e.g. Dewees, 1995; Drechsel et al., 1996). The absolute value of the NPV for an agroforestry option thus provides much information about the private profitability of the option, over its life span. The quality of the information is increased when a range of discount rates is used, when NPVs between different options are compared and when sensitivity analysis is undertaken. The concept must be further interpreted and analysed in order to also provide information about the adoptability of an agroforestry option, in addition to its private profitability.


29

Interpretation of results from private cost–benefit analyses Since NPV is an aggregate estimate of profitability over time, it is essential to identify the length of time during which net losses will be incurred by adopters before NPV starts becoming positive. This is called the break-even point of an agroforestry option. The evidence reported in the literature indicates that many agroforestry practices have break-even points longer than 1 year. An example concerns the adoption of trees on contour lines in Java. It takes a minimum of 5 years for the cumulative on-farm benefits to exceed the initial cost outlays in these agroecosystems (Barbier, 1990). In the example of adoption of improved fallows in western Kenya, it takes 4 years for this to happen (Swinkels and Franzel, 1997). In their review of multiple agroforestry practices in several countries, Current et al. (1995) found break-even points ranging from just under 2 years (for alley cropping) to more than 9 years (for woodlots). Adoption of agroforestry

Total benefits, total costs of agroforestry practice

TBa

TBb

TC3 = TBa3

TC

TBb3

TBa = annual total benefits, scenario a TBb = annual total benefits, scenario b TC = annual total costs

0 Year 1

Year 3

Year 10

Time

Fig. 2.3. Farm-scale costs and benefits of agroforestry adoption over different time periods.

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options that entail losses during 2 to 5 years (these are the most common break-even points reported in the literature) thus requires that farmers have the financial ability to absorb these losses. Clearly, not all resourcepoor farmers in tropical countries will have such an ability. The soil fertility management and agroforestry practices that have the highest likelihood of being adopted by these farmers are those that result in sufficiently significant short-term benefits to compensate for the costs of implementation. This is summarized in Fig. 2.3, in which these costs of implementation are shown as a decreasing function over time. The corresponding benefits curve however, increases exponentially with time, for the reasons mentioned above. Two scenarios are represented in Fig. 2.3. In scenario a, total benefits (TBa) increase rapidly over time whereas in scenario b they increase relatively slowly. If the planning horizon of the farmer is up to 3 years and if scenario a prevails, then adoption of agroforestry practices can be expected. If scenario b prevails and the farmers’ planning horizon is around 3 years, then adoption is unfeasible because, over this time horizon, total benefits (TBb3) remain below total costs. This balancing of monetary and non-monetary, present and future, on-farm costs and benefits over the duration of the farmer’s planning horizon is a fundamental economic process affecting soil fertility management at the farming system scale. The few economic analyses of agroforestry practices for soil fertility management that rely on actual on-farm data provide an illustration of this when they stress that a constraint to adoption is that farmers often do not have sufficient capital to be able to withstand the few years of low, or even negative, profits which precede the years of relatively high profitability (e.g. Scherr, 1995; Grist et al., 1999). It is enlightening to note that even in the economic context of the USA, where farmers receive government subsidies (called cost-sharing), tax credits and deductions for adopting agroforestry practices, rates of adoption of agroforestry practices have been lower than expected for essentially the same reasons. A survey of farmers in the southern USA revealed that, although these farmers were aware of the environmental benefits of agroforestry, ‘considerable uncertainty remained’ about the potential profitability of various agroforestry systems in the minds of these farmers (Zinkhan and Mercer, 1997). In summary, the fundamental economic processes that occur at the farm scale are as follows. •

•

Farmers bear all the costs of adoption of improved agroforestry and soil fertility management practices, and these costs start occurring from inception of adoption and continue over time. Farmers accrue the monetary benefits of adoption and some of its nonmonetary benefits.


• •

•

31

These benefits are cumulative over time and tend to be relatively low during the initial years following adoption. The time lag between costs and benefits of adoption for farmers, coupled with the fact that small-scale resource-poor farmers have short-term planning horizons and high discount rates, implies that rates of adoption can be very disappointing to scientists and policy makers. A major implication for scientists is that it is essential for them to design practices which minimize up-front costs of adoption (labour, capital and land).

The simplest and most effective economic method available for quantifying these processes for specific agroforestry practices is private cost–benefit analysis. It encompasses the computation of the NPV of a practice, from the perspective of an individual farmer. Very useful insights into the probability of adoption of a practice can be gained by: (i) comparing the NPV of an agroforestry option with the NPV of other land-use options of relevance to farmers; (ii) undertaking a sensitivity analysis of NPV; and (iii) estimating the break-even point of an agroforestry option.

Other economic methods: modelling Once the above processes are quantified, there are a few additional methods that can be used for simulating the profitability, adoptability and biophysical consequences of agroforestry practices. These methods all entail some form of modelling of the economic processes just discussed, in relation either to biophysical processes or to various social and cultural variables. Most of them have been developed very recently and have been validated in only one set of circumstances. They are mentioned here only for reference purposes as they lie outside the scope of this book. An example of a bioeconomic model, that is, a model that integrates economic and biophysical variables and processes, is provided by Grist et al. (1999), who used SCUAF, an agroforestry biophysical model, in combination with economic spreadsheets. Another example can be found in Shepherd and Soule (1998), who have built a farm simulation model which simulates the effects of different land-use management strategies, including agroforestry options, on soil processes and nutrient-limited plant production and on profitability. Adesina et al. (2000) provide an example of a model (Logit) that explains actual farmers’ decisions to adopt alley farming on the basis of a number of explanatory variables. These variables reflect the land tenure system, the socioeconomic characteristics of farmers (e.g. level of education, membership in farmers’ association) and village-specific

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characteristics (e.g. land pressure, erosion index and importance of livestock in the village).

2.5 Landscape and Global Scales: Soil Fertility and Agroforestry Trees as Part of Natural Capital It was shown in Section 2.3 that farmers’ decisions at the farm scale not only are shaped by variables determined at higher levels than the farm in the hierarchy of systems but also have consequences at these higher levels. Two concepts from ecological economics are particularly useful in understanding these interactions between scales. These are the concepts of natural capital and of ecosystem services. It has been demonstrated that soil nutrients and trees are part of natural capital (Izac, 1997, for soil nutrients; Izac and Sanchez, 2001, for agroforestry trees). Natural capital comprises all the natural resources that provide useful goods and services for mankind. More specifically, natural capital is defined as stocks of resources generated by natural biogeochemical processes and solar energy that yield useful flows of services and amenities for the present and into the future (Izac, 1997). Natural capital generates ecosystem services which are the processes ensuring the productivity, integrity, maintenance and resilience of ecosystems. The ecosystem services generated by soil nutrients include nutrient cycles, soil fertility, plant nutrition and carbon sequestration. The ecosystem services generated by agroforestry trees include, for instance, erosion control, water cycling, pest and disease control, and biodiversity. An important characteristic of ecosystem services is that they occur over different spatial and temporal scales (Izac, 1997). Carbon cycling, for instance, is a global cycle related to climate change, taking place over the atmosphere, soils and ocean-bottom sediments, and doing so over relatively long periods of time. At the plot scale, carbon is linked to soil fertility. Table 2.3 illustrates the principal ecosystem services of trees at different spatial scales. Since soil nutrients and trees (natural capital) generate different ecosystem services at different spatial scales, different members of society will be affected by these ecosystem services. Likewise, the management of these ecosystem services and changes in this management affect these different groups in society. When the non-monetary benefits of soil fertility management through agroforestry were discussed, it was pointed out that farmers may be quite indifferent to some of these benefits or ecosystem services (e.g. soil biodiversity and carbon sequestration). These benefits are, however, valued by other members of society. National societies in tropical countries do value sustainability of food production and


33

Table 2.3. Principal ecosystem services of agroforestry trees at different scales. Scale

Ecosystem functions

Farm

Food production Nutrient cycling Erosion control Water cycling Genetic diversity Microclimate regulation

Watershed/village/landscape

Decreased poverty Erosion and sedimentation control Water cycling Refugia, pollination, biological control (landscape patches)

Region

Decreased poverty Decreased deforestation and desertification Biodiversity Water cycle

Global/supraregional

Greenhouse gas regulation Climate regulation Biodiversity Rural poverty alleviation

biodiversity, for instance, since most countries have food security and biodiversity strategies and the global society does value carbon sequestration, as demonstrated by the fact that there is an international market for sequestered carbon. From an economic perspective, such benefits of the ecosystem services of agroforestry systems for soil fertility management are positive environmental externalities. These are defined as follows. Investments by farmers in agroforestry systems trigger flows of non-monetized benefits accruing to different groups in society, called environmental externalities. Farmers accrue some of these benefits, largely those related to increased risk buffering capacity and sustainable food production (in addition to accruing the monetary benefits of agroforestry). National society will receive the benefits of decreased rural poverty, watershed protection, increased biodiversity, more sustainable agricultural production and increased food security. Global society will enjoy increased carbon sequestration and biodiversity benefits. These various environmental externalities are likely to be highly valued by national and global societies because these have a longer planning horizon than individual farmers. The relative significance of

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ecosystem functions vis-à-vis food and income production is difficult to appraise. An attempt at evaluating these functions has, however, been made. Costanza et al. (1997) estimated that the global value of 17 ecosystem services for 16 key biomes is about US$33 trillion per year. This is excluding the worth of the stocks of natural capital in these biomes, as stocks of natural capital are too difficult to evaluate from an economic perspective. By comparison, the world gross national product (GNP) is only worth some US$18 trillion per year and the food produced by all croplands amounts to about US$0.13 trillion per year (Costanza et al., 1997). These estimates give an indication of the order of magnitude of the value to society of the ecosystem services generated by natural capital compared to the value of agricultural production from croplands and the value of total economic production, or GNP. The economic method most appropriate for assessing the various environmental externalities associated with agroforestry practices is social and environmental cost–benefit analysis. This consists of a complex set of economic methods (sometimes called shadow pricing) for assessing all nonmonetary benefits and costs for different groups in society, in addition to the assessment of the private costs and benefits described in the previous section (see Mishan, 1976, for details). Furthermore, it requires that all price distortions due to government or policy failures be redressed as the analysis is conducted from the perspective of society as a whole (Mishan, 1976, provides the best explanation of these various methods, as well as of their shortcomings). This is expressed as: NPV =

n

Â t =1

( A + I ) – (D + S ) (1 + r ) t

t

t

t

t

(2.4)

s

where: rs = social rate of discount; At = vector of all monetary benefits of an agroforestry option during year t, after adjusting prices for relevant market failures; It = vector of all non-monetized benefits received by different groups in society during year t, after adjusting prices for relevant market failures; Dt = vector of all monetary costs of an option, after price adjustments for relevant market failures, in year t; St = vector of all nonmonetary costs of an option, after price adjustments for relevant market failures, in year t. The social rate of discount (rs) is the rate at which society is willing to trade present benefits for future benefits. Since society has a longer time horizon than individuals, this rate is generally significantly lower (e.g. 5%) than the individual rate of discount of relevance to farmers in tropical countries. To date, social cost–benefit analysis has not been applied to the assessment of agroforestry options, probably because there are some currently unresolved methodological challenges. This is, therefore, an area


35

for future research. The International Centre for Research in Agroforestry, for instance, initiated in 2001 a global research project the objective of which is to assess, from an economic perspective, the various environmental externalities of key agroforestry practices. The ecosystem services and environmental externalities associated with agroforestry systems and soil nutrients indicate that what is an optimal level of adoption of agroforestry practices from the viewpoint of farmers is a suboptimal level of adoption from the perspective of national and global society. This situation is represented in Fig. 2.4. The marginal (monetary and opportunity) costs of adoption of agroforestry practices are compared with the marginal (monetary and non-monetary) benefits of adoption received by individual farmers, those received by the national society and, finally, those received by the global society. Marginal costs of adoption are incurred uniquely by farmers, in the absence of any policy geared at implementing a cost-sharing scheme. Marginal benefits of agroforestry are higher for the national society than for individual farmers, and yet higher for the global society, for the reasons just mentioned. Figure 2.4 shows that the level of adoption of agroforestry practices that is optimal at the farm scale (from the viewpoint of farmers) is QP, whereas the level of adoption national society regards as necessary is QN, and QG is the globally optimal adoption level. In the absence of any policy, farmers will adopt level QP, which is suboptimal from society’s viewpoint Marginal cost (individual)

Marginal benefits (global society)

Marginal costs and benefits of agroforestry adoption

Marginal benefits (national society) Marginal benefits (individual)

0

QP

QN

QG

Level of agroforestry adoption

Fig. 2.4. Regional and global level trade-offs between the costs and benefits of agroforestry systems at a given point in time.

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because it does not take into account most of the ecosystem services of agroforestry systems. These ecosystem services are therefore unlikely to be fully realized. These trade-offs between the individual and societal costs and benefits of agroforestry adoption (represented in Fig. 2.4) are fundamental economic processes at the regional and global scales of analysis. The economic processes discussed in this chapter indicate that private and social interests concerning soil fertility management and agroforestry in tropical countries do not coincide. The extent of agroforestry practices voluntarily adopted by farmers (that which is in their own best interest) will almost certainly be inferior to that which is socially optimal. It will not be optimal or effective or equitable to expect resource-poor small-scale farmers in tropical countries to bear the full costs of adoption while national and global societies receive significant benefits from this adoption. In such circumstances, providing farmers with information and advice on soil fertility management and agroforestry practices will definitely be insufficient for bringing about socially optimal adoption levels. Some policy measures will be needed for social optimality to occur. Discussions of possible policy instruments, from cost-sharing schemes, to communitybased management and carbon offset mechanisms can be found in the literature and lie outside the scope of this chapter (see, for instance, Babu et al., 1995; Dewees, 1995; Adesina and Coulibaly, 1998; Donovan and Casey, 1998). The point we wish to emphasize here is that scientists working on the design of soil fertility management practices have a moral responsibility to ensure that their results are communicated to policy makers in the clearest and most effective manner possible. Sufficient information is now available about relevant economic processes at play when decisions to adopt are made by farmers, that the scientific community can no longer continue to argue that all that is needed for optimal levels of adoption to take place is good extension work and rational farmers. We now know that farmers are indeed rational and that the best extension system in the world is not set up to develop the policy instruments needed to bridge the gap between individual and societal benefits and between individual costs and societal benefits. In summary, the key economic processes at the landscape and global scales are as follows. • •

Agroforestry trees and soil nutrients are part of natural capital. As such, they generate ecosystem services at different spatial scales. These ecosystem services are positive environmental externalities. As such, they are non-monetized benefits of adoption that occur off farm and accrue to different groups in society (national and global society).


• •

• •

•

37

These environmental externalities have a high value to national and global societies. Farmers do not take many of these environmental externalities into consideration in their decisions to adopt because these are off-farm benefits. Consequently, optimal levels of adoption from the farmer’s perspective are lower than the socially optimal levels. Social and environmental cost–benefit analysis is an appropriate method for assessing these different processes. This is a new area of research in agroforestry. It is essential for scientists to communicate the results of their work to policy makers to increase levels of adoption towards socially optimal levels.

Chapter 3 Designing Experiments and Analysing Data R. COE,1 B. HUWE2

AND

G. SCHROTH3

1World

Agroforestry Centre, International Centre for Research in Agroforestry (ICRAF), PO Box 30677, Nairobi, Kenya; 2Institute of Soil Science and Soil Geography, University of Bayreuth, 95440 Bayreuth, Germany; 3Biological Dynamics of Forestry Fragments Project, National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil

3.1 Synopsis Appropriate experimental design and analysis of data are fundamental to furthering our understanding of the impact of trees on soil fertility. Although there are many standard statistical texts that describe procedures for agronomic or forestry trials, the inclusion of one or more tree components in an agricultural context tends to increase both the spatial variability and the time horizon that have to be considered. This chapter makes reference to places where standard designs and analyses are described, but focuses on aspects of particular importance in an agroforestry context. Given the long-term nature of many agroforestry practices, setting appropriate objectives for individual experiments is of vital importance but is seldom discussed in standard texts and is often the root cause of failure to achieve useful outcomes from agroforestry trials. Here, advice on setting objectives is given before consideration of appropriate treatment structures, layout and replication. Experiments with tree fallows are given special attention. They have become particularly important in soil fertility research over the last decade and require careful consideration because they generally involve a large number of factors, are subject to interference among plots and include treatments with different fallow lengths and hence timing and duration. The spatial and temporal heterogeneity in soils induced by the presence of a relatively few, large plants that grow over several years complicate sampling strategies, so advice is given on the most efficient sampling schemes to use to meet different types of experimental objectives. © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

39

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Collection of data is of little value unless they are appropriately analysed, but objectives for analysis are often modified as research proceeds, so a process of exploratory analysis followed by application of formal statistical tests is recommended. Particular advice is given on simple methods of dealing with repeated measurements in time and space and the appropriate statistical software to use in data analysis. The final part of the chapter deals in some detail with methods for analysis of spatial structure. This is likely to be of increasing importance in agroforestry research, where spatial heterogeneity at various scales is increasingly embraced as an important component of agroforestry practices and landscapes, rather than being seen as something that has to be minimized. The techniques for spatial analyses are relatively new and have not yet been widely applied in agroforestry contexts. They are discussed here in sufficient detail for readers to decide how to approach analysis of spatial structure and make reference to the most appropriate literature where further details and examples are available.

3.2 Experimental Objectives, Treatments and Layout (R. Coe) The design and analysis of experiments for investigating fertility effects of agroforestry follow the principles and practices that have been developed for field experimentation over the last century or more. These principles are summarized here, with references given to more complete descriptions. Checklists of points to remember when designing an experiment may be useful and are given elsewhere by Coe (1997a, 1999). This chapter focuses on some aspects of experiments for investigating impacts of trees on soil fertility in which it can be difficult to apply these well-established principles.

Identifying the objectives of an experiment The objectives of an experiment determine every aspect of its design, so are a key to a sound design. The objectives must be: • • •

•

Clear: objectives stated vaguely or ambiguously lead to confusion later; Complete: incompletely determined objectives mean that some design decisions cannot be made; Relevant: applied research has aims of solving specific problems and each experiment must have a clear place in the problem-solving strategy; Capable of being met by an experiment: not every research objective can be met by an experiment.

Designing Experiments and Analysing Data

41

Much of the rest of this book is aimed at helping researchers determine what areas of research might be fruitful and how to go about making appropriate measurements of key variables. However, this is not the same as identifying objectives for individual trials that meet the criteria above and which will allow all decisions about the design to be made. Few of the texts on experimental design have much to offer on the problem of statement of objectives, even though the problems scientists encounter in many field trials originate from objectives that are confused or contradictory. The classic book by Cochran and Cox (1957) devotes half a page to the topic and more recent books do no better, maybe because there is little structured advice that can be given. A recent exception is Robinson (2000). There are, however, some common problems which can be avoided. First, objectives as vague as ‘to see what happens’ cannot be used to design a trial. If the research is still at the stage of having no clear hypotheses or quantities to measure then an experiment is not (yet) needed. Secondly, careful reading of objectives often reveals them to say nothing more than ‘the objective of the experiment is to compare treatments’. Objectives should be expressed as knowledge gaps to fill, that is, hypotheses to test or quantities to estimate, which lead to a definition of treatments, rather than starting with the treatments themselves. Most experiments have multiple objectives but these have to be mutually consistent. A common conflict is between objectives with a technology perspective, aimed at evaluating options for farmers, and those aimed at understanding processes. These two perspectives can often lead to different requirements of treatments and management. For example, Rao et al. (1998) describe a trial in which Senna spectabilis was deliberately used, rather than a species of greater importance to farmers in the region, because the roots were easily distinguished from those of the accompanying crops. In technology development trials, a further common difficulty is simply that of choosing where to start among the many aspects that could be studied. Cooper (1997) describes an approach to thinking through this, based on identification of critical constraints, and Franzel et al. (2001) discuss the appropriateness of different types of on-farm trials, with varying degrees of farmer and researcher control, for meeting different objectives.

Defining experimental treatments The key ideas needed to define treatments for an experiment are as follows. • Contrasts: most objectives can be reduced to the need to compare two or more treatments, either to detect whether a difference exists or to measure the size of the difference. Clearly the treatments to be

42

•

•

•

R. Coe et al.

compared or contrasted determine the treatments required in the experiment. Controls: control treatments are nothing special, simply the standards against which other treatments are compared. They do not have to be defined as ‘zero input’, ‘farmer’s practice’ or similar, but will be chosen depending on the objectives. Factorial treatment structure: this can arise naturally from the hypotheses or objectives, but can also be used to test several unrelated hypotheses within one trial more efficiently than with separate trials. Quantitative levels: when treatments involve different quantities of something, then the actual levels required may not be defined by the objectives, but guidelines on choosing them are available.

Mead (1988) gives a thorough practical discussion of statistical issues in choice of treatments. Defining suitable control treatments should not be difficult if the objectives are clear, but there can be conflicts if the trial has multiple objectives. Commonly, farming system and biophysical objectives conflict and in some cases (at least) two controls are needed. Examples are a biophysical control for process-oriented research and a farming system control for applied studies, or a forestry and an agricultural control because both of these represent well established alternative land uses (Dupraz, 1999). The concept of controls is ill-defined and it is clearer to simply consider the contrasts needed to meet the objectives and then the treatments needed for the defined contrasts. Quantitative treatments are common in experiments concerning agroforestry and soil fertility – for example, amounts of organic or inorganic input and density of trees. For these types of treatments the objective usually reduces to estimation of a response curve. Choosing treatments involves choosing the range of levels (minimum and maximum), the number of different levels, the spacing between the levels and the degree of replication of each level. There is a mathematical theory of design applicable where a smooth response curve is expected, from which the following general rules emerge. • •

•

Make the range as wide as feasible. Make sure the highest levels are well past any optimum. Limit the number of different levels; estimating a straight line needs just two. A third will allow its straightness to be checked. More complex curves may need four or five levels but it is very rare to find an example that needs more different levels than this. Put levels closest together where you think the response will change fastest. In the absence of any other information use equally spaced levels.


43

When there are two or more quantitative treatments there are important alternatives to complete factorial sets of treatments. Many agroforestry experiments involve treatments of distinct agricultural systems, not just relatively minor variations of the same system. This means it can be difficult to define the management of the differing plots in a way which does not lead to confounding of treatment effects with management variables. A simple example from Kenya involved screening seven tree species for their effect on intercropped maize. An eighth treatment of monocrop maize was included. As this was a phosphorusdeficient site, trees were given a starter dose of phosphorus fertilizer but the monocrop was not, resulting in confounding of effects of trees and phosphorus fertilization. The design is acceptable for comparison of systems, to be assessed perhaps on the basis of profitability, but biophysical data on, for example, phosphorus cycling or even crop yield are difficult to interpret. Dupraz (1999) gives other examples of this problem, including confounding effects of pest and weed control.

Experimental layout Layout of a trial involves choice of locations, definition of the plots, characterization of the site, decisions on replication, choice of blocks and allocation of treatments to plots. Much of the theory and practice of all these developed for agricultural trials is relevant to agroforestry trials. There are many books on the topic. Dyke (1988) gives practical field advice. Mead (1988) covers the more statistical theory. Notes by Coe (1997b,c) and Stern and Adam (1997) are also relevant. There are two important differences between typical soil fertility experiments involving annual crops and those with agroforestry treatments which arise because of the differing time and spatial scales involved. •

Trees may take several years to produce the effects of interest, so trials will usually be in the ground much longer than those with annual crops. If a trial of one growing season turns out to have been poorly designed or laid out then it can be repeated the following season. This is not true for an agroforestry trial in which design faults associated with the tree component may not be apparent until years after it was planted. Hence, it is worth investing resources in ensuring the layout is as sound as possible. In particular it is worth characterizing the site to identify heterogeneity, which can then be allowed for by a combination of blocking, avoidance and increased replication (see also Section 3.6). Another implication of the long time horizon of agroforestry experiments is that they will inevitably be used for

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purposes other than those for which they were originally planned. It is therefore worth building some flexibility into the design. This means keeping treatments simple, using large plots that can be split later and including extra replicates that may be modified as needed (for example, by destructive harvest). Trees are typically larger than annual crops, which also has implications for the layout of experiments. Most important is the avoidance of interference between neighbouring plots. Above-ground interference occurs through such effects as shading, windbreak effects and lateral transport of tree litter by wind. These may not be easy to control but at least can be seen and perhaps avoided. Below-ground interference occurs as tree roots extend beyond their nominal plot and capture resources from surrounding areas, either from other plots or from outside the experiment. In many trials it is clear that this has resulted in biased treatment comparisons. If a plot with trees is adjacent to a crop-only plot then the tree plot may capture extra resources and the crop plot may be deprived of resources relative to a situation in which both are grown in very large plots, resulting in a biased treatment comparison. The options for avoiding the bias are either to leave very large border areas or to physically prevent interference.

It is difficult to judge the size of borders required in any situation; this clearly depends on the characteristics of the species involved, the site and the length of time the trees will grow. Van Noordwijk et al. (1996) discuss

Fig. 3.1. Lateral root from a Sesbania sesban fallow plot (on the right) grown into a crop plot (Cajanus cajan) in about 4 months.


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factors affecting tree root growth in agroforestry systems and make the point that it is very difficult to give general guidelines for the lateral extent of roots. Certainly, in dry areas, extensions of over 40 m have been recorded. Figure 3.1 shows a lateral root of Sesbania sesban known to have extended over 6 m in 4 months. Physical prevention means installation of below-ground barriers or regular root pruning between plots. Barriers are of limited value as roots can quickly grow under them and up into adjacent plots (Rao and Govidarajan, 1996). Pruning below ground is expensive as trenches have to be re-opened each time it is done, possibly four times a year. Trenches cannot usually be left unfilled between prunings because of their impact on water movement and the likelihood that they will behave similarly to solid barriers, with roots growing under them. A third alternative that has been suggested is to lay out trials so that plots are not adjacent. This will probably decrease the precision of the experiment but, more importantly, will not remove the small plot bias as trees will still be capturing resources from outside their nominal area. In fallow experiments it is common to use large plots with large borders for the fallow phase of an experiment, then to split the plots for further treatment factors during the cropping phase. Remember that the original borders may still have to be left as they may be untypical even after the trees have been removed. An alternative approach recognizes that, within many farming systems, agroforestry will be used in small patches in a landscape. This landscape will be full of edges between agroforestry and other land uses, and the lateral interactions between them may be an important component of the way the system functions. Indeed, this agricultural landscape mosaic with trees has been defined as agroforestry (Leakey, 1996). The implication for research is that interactions should not be removed from trials but incorporated into the objectives (van Noordwijk, 1999). Designing trials with a system focus using this notion is feasible, though challenging. The problems are likely to be both in selecting useful objectives from the myriad of possible topics, and installing, managing and monitoring trials of the size required. The alternative is to measure lateral flows in simpler experiments and then attempt to model the landscape level effects of agroforestry systems. The longer time scale and larger plots needed in typical agroforestry trials, compared with annual crops, can make trials with farmers difficult. Farmers have to commit relatively large areas of land for long periods. The areas required mean that often each farmer can only test a few treatments and the ideas of incomplete blocks are needed to find effective designs. The long time periods also mean that both drop-outs and modifications by farmers are common, resulting in more replicates being needed.

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Number of replicates The number of replicates required in an experiment is straightforward to estimate in theory, but can be difficult in practice as information required is unknown before the trial starts. Replication serves three purposes in an experiment: • •

•

to allow estimation of the random variation, and hence of the precision of contrasts of interest; to increase the precision of estimating important quantities, the average of a larger sample being more precise than that of a smaller one; and as insurance against disasters that lead to loss of parts of the trial.

The number of replicates needed to satisfy each of these purposes has to be looked at. The quality of the estimate of a variance depends on the degrees of freedom associated with it. For a simple randomized block design the variance relevant to estimation of treatment differences is the residual or error variance. The degrees of freedom in the residual line of the analysis of variance table indicates how well that variance will be estimated from the design. Experimenters should aim for residual degrees of freedom to be at least ten, though one or two less than this may be acceptable. Residual degrees of freedom less than six are hopeless. For more complex designs which have several different layers, such as split-plot or multisite designs, there is more than one variance relevant to estimation of treatment effects, and each has to have sufficient degrees of freedom. The researcher in these circumstances has to be able to produce an outline analysis of variance for the design and check that degrees of freedom for each relevant error line are adequate. Some software can produce these outline analyses, but any software can be forced to do so by putting in dummy data, as the degrees of freedom depend only on the design, not on the data values. In agroforestry trials it is not unusual for the degrees of freedom to be different for different variables. For example, a trial that compares monocrop maize with maize intercropped with trees has more plots with crop data than with tree data. The analyses of variance for tree and crop variables will, therefore, have different degrees of freedom, so both need checking. Estimation of replication required for the second purpose, controlling precision, requires experimenters to have an idea of: (i) the size of relevant error variances likely to occur in their trial, and (ii) the precision they actually require. The first of these has to be estimated from previous experience with other trials of a similar type in a similar environment. Although every trial should be different from all predecessors, there are few experiments that are so novel that it is not possible to get some idea of the expected levels of variation. The variation may be expressed as a


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variance (s2) or independently of the units of measurement as a coefficient of variation (cv), which is simply the standard deviation divided by the mean. The precision required is most usefully expressed as the standard error or width (w) of confidence interval needed for critical quantities. Alternatively, it may be expressed as the size of effect that the trial needs to detect as significant. Formulae for the number of replicates may then be deduced. For example, for an approximate 95% confidence interval of width w for the difference between two means in a randomized block design, the required number of replicates (r) is r = 32

s2 w2

(3.1)

Software is available for producing formulae for other designs and other specifications of the problem (Thomas and Krebs, 1997). For the third reason for replicating, insurance, it is harder to judge the number required. If an experiment has a short lifespan (say one season) then there is little justification for insurance, unless perhaps under farmer management where there is some uncertainty about the likelihood of individual farmers adhering to important aspects of a protocol. However, if a long-term trial is in a situation where it might be at risk (say from fire) then inclusion of a few extra replicates may be judged worthwhile, particularly given the other reasons for building some redundancy into the initial design, mentioned above. The situation for the simplest on-farm trials, designed to estimate the mean difference between two treatments, is no different from that for onstation trials, and the same information is required to estimate sample sizes. Variation at all levels tends to be higher, so numbers of replicates needed are often large. The large plots needed in many agroforestry trials mean that it is often only feasible to have one replicate per farm, and farms should be treated as blocks. If it is only possible to have less than one replicate per farm then incomplete block designs are useful, with an implication for the number of replicates needed. The statistical principles are described well by Mead (1988), and software is available to help find suitable designs. Many on-farm trials have more complex objectives than simply comparing two treatment means, but are concerned with performance of agroforestry in different farm and landscape situations. Very often there is a hierarchy of levels in the design from individual plants at the lowest level to communities or watersheds at a higher level. At each of these levels there are design questions to resolve concerning the number of units, the selection of those units and the treatments or interventions to be compared on those units (Table 3.1). The same ideas are needed for determining the number of replicates in these more complex designs as in simpler ones.

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Table 3.1. Hierarchy of levels and questions to be answered in design of on-farm experiments. How many?

Layer

Which ones?

Who decides?

What intervention?

What is measured?

Village Landscape position Farm Niche Plot Tree

3.3 Fallow Experiments (R. Coe) The principles underlying the design and layout of fallow experiments are no different from those for other experiments, the key to a sound design being very clear and exact objectives for the trial. However, fallow experiments can give rise to particular difficulties because of the potentially large number of factors to investigate, the potential for plot to plot interference and the fact that time may be involved as a treatment. If the experiment is system focused, aimed at evaluating the effect of management factors, there are questions concerning fallow establishment, management, clearing, post-fallow cropping and starting a second fallow cycle. Techniques similar to those described in Tripp and Wooley (1989) can be used to prioritize factors for investigation. The idea is to choose criteria such as ease and cost of doing the research, the extent to which the factor is critical to adoption of the technology, the length of time the research will take and the chance that it is successful. The various factors that could be investigated are then scored on each of the criteria to produce a ranking. The number of treatment combinations can sometimes be reduced by assumptions such as, for a given location and species, that

Fertility

Fallow

Crop

Time

Fig. 3.2. Schema of soil fertility development during cropping and fallow cycles.


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Table 3.2. Design of a fallow experiment comparing fallow lengths of one, two and three seasons. Season Treatment

1

2

3

4

1 2 3

F F F

C F F

C F

C

F, fallow; C, crop.

there is a direct relationship between the fertility effect of the fallow and the fallow biomass (Mafongoya and Dzowela, 1999). When the trial has a process focus, the process objectives will determine treatments. Note in particular that species for fallows can often be chosen in a way that gives the trial much more value than a simple comparison of a range of likely species. An example is the comparison of four species deliberately chosen to evaluate the two characteristics of deep-rooted vs. shallow-rooted and nitrogen fixing vs. non-fixing. The control treatments might be a natural fallow, continuous cropping, a herbaceous fallow or even bare ground (Hartemink et al., 1996), the choice, of course, depending on the exact objectives (see Section 3.2). Much applied research on fallows is concerned with factors associated with the rate at which they restore fertility and the rate at which crops reduce it – the shape and slopes of the curves in Fig. 3.2. Investigation of fertility recovery during the fallow phase requires comparison of fallows of different lengths. A design to look at fallows of one, two and three seasons may involve treatments as shown in Table 3.2. The crops in seasons 2, 3 and 4 will reflect the changes in soil fertility following one, two and three seasons of fallow. However, the expected pattern (Fig. 3.2) will be confounded with the season-to-season fluctuations in weather and disease patterns. For this reason the phased entry design of Table 3.3 is preferable. Table 3.3. Design of a fallow experiment comparing fallow lengths of one, two and three seasons (version 2). Season Treatment

1

2

3

4

1 2 3

* * F

* F F

F F F

C C C

F, fallow; C, crop.

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Now the effects of differing fallow lengths are assessed by comparison within a single season. However, the design raises the two following questions. •

•

What should be put in the cells marked * ? At the start of the first season the whole site is presumably in the state required for the experiment. To avoid confounding fallow length with other factors we need the soil in treatment 2 plots to be the same as this at the start of the second season, and those of treatment 1 plots at the start of the third season. There is no practice that can guarantee this. If the trial is concerned with farming systems then it is common for the initial condition to be that state of degradation in which a farmer would abandon cropping and start a fallow phase. It is often assumed that one or two further seasons of cropping on these degraded plots will not substantially change the soil further, so the design used is as in Table 3.4. The designs in Tables 3.3 and 3.4 are appropriate if the seasonal effects (such as weather) apply to the crop but not the fallow. Although it is likely that in general a crop will be more sensitive to seasonal differences than a fallow, this need not be the case. Suppose the third season was a very good one for fallow establishment, but that the second season was not. This difference may be reflected in the following crops and be confounded with the comparison between the two seasons. A way around this is to repeat the sequences in more than one season (Table 3.5). It is clear that this design can quickly grow to something unmanageable, both because of the number of plots and the time involved. Of course there is no guarantee that the seasons sampled with just one repeat of the sequences will be sufficient. Sites are sometimes substituted for seasons, putting treatments 1, 2 and 3 down simultaneously in a number of locations.

When looking at the cropping phase of the sequence there are similar considerations. If we want to measure fertility decline over, say, three Table 3.4. Design of a fallow experiment comparing fallow lengths of one, two and three seasons (version 3). Season Treatment

0

1

2

3

4

1 2 3

D D D

D D F

D F F

F F F

C C C

F, fallow; C, crop; D, crop on degraded soil.


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Table 3.5. Design of a fallow experiment comparing fallow lengths of one, two and three seasons, repeated through two different seasons. Season Treatment

0

1

2

3

4

5

1 2 3 4 5 6

D D D D D D

D D F D D D

D F F D D F

F F F D F F

C C C F F F

C C C


seasons of cropping following a fallow, the comparisons require a design as in Table 3.6. In the fourth season the first, second and third crops following a 1 year fallow are compared. It is clear that the requirements for comparing differing cropping and fallow periods are mutually incompatible. It is not possible to both start and end different treatment sequences at the same time if their lengths differ. For this reason compromise is necessary, the usual solution being to hope that fallows are less sensitive to seasonal variation than crops. The fallow plots can be measured to find out whether growth is in fact similar in different seasons (survival and biomass at the end of the first season of growth would be appropriate for this purpose), leaving open the question of what to do if this turns out not to be so. A typical design to study the decline in fertility following fallows with durations of one and two seasons uses treatments 1–6 in Table 3.7. An alternative approach uses the idea that it is possible to define a control treatment (perhaps fully fertilized) that responds to weather and other season-specific factors but is constant with respect to soil fertility. A trial to look at decline in fertility following fallow might then use just treatments 3, 6 and 7 from Table 3.7. Crop yield of treatments 3 and 6 Table 3.6. Design of a fallow experiment comparing cropping cycles of one, two and three seasons. Season Treatment

0

1

2

3

4

1 2 3

D D D

D D F

D F C

F C C

C C C


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Table 3.7. Design of a fallow experiment comparing cropping cycles of one, two and three seasons following one or two seasons of fallowing. Season Treatment

0

1

2

3

4

5

1 2 3 4 5 6 7

D D D D D D D

D D D D D F Cont

D D F D F F Cont

D F C F F C Cont

F C C F C C Cont

C C C C C C Cont

F, fallow; C, crop; D, crop on degraded soil; Cont, control.

during seasons 3, 4 and 5 would be assessed by measuring them relative to those in treatment 7. The approach depends on the assumption (untestable with the design) that there is a scale on which there is no treatment ¥ season interaction, which seems unlikely, particularly when considering variables other than crop yield. Note that if treatments 1 to 7 of Table 3.7 are used then the value of this approach can be assessed by comparing the trends derived from the time series with those measured in season 5.

3.4 Measurements and Sampling Designs (G. Schroth, R. Coe) Much of this book is concerned with measurement and there are many other sources with practical information both on what to measure and how to measure it. When planning experiments, it is worth thinking of the following three categories of measurements. •

• •

Measurements that are needed to characterize the site in which the trial takes place, but do not vary between treatments. These include, for example, soil type and rainfall. Responses specified by the objectives, which might be soil nitrogen and plant growth. Measurements needed to understand variability in the key responses, which might be soil depth or texture and are often forgotten. They can be specified once the nature of the variability becomes apparent and field observation suggests some possible causes.

A special case of the first type of measurements are those that are specifically made to show that basic site characteristics do not vary significantly between treatments. This is especially important when proper


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replication of treatments is not possible, because an experiment has not been established as such, but is rather a comparison of plots that have been under different vegetation cover or land use for a certain time (such as comparing agroforestry, conventional agriculture and natural forest). Such ‘unplanned’ comparisons often provide information that planned experiments cannot yield for practical reasons, especially if long time periods under a certain land use are needed. However, they always suffer from the difficulty of establishing whether the initial soil conditions were similar among sites. Sanchez (1987) suggested particle size distribution in the profile as the principal criterion for checking the comparability of sites. Nearly all measurements in experiments require sampling of some sort and strategies for sampling soils are discussed in detail here. There are several recent descriptions of soil sampling procedures for agricultural (James and Wells, 1990) and environmental purposes (Crépin and Johnson, 1993). When adapting such recommendations for agroforestry situations, it is essential to be sure about the aim of the study, as this will affect decisions about the sampling plan as well as sampling depth and timing. There are two key aspects to be clear about. •

•

The level at which replication is needed to estimate precision. As an example consider plots of an experiment to be sampled to determine differences in soil properties under different treatments. Although several samples may be taken in each plot, it is not usually necessary to estimate the precision of values for each plot. Instead we need values from replicate plots to estimate precision of differences between treatment means. Whether it is necessary to compare different parts of the plot or field. In many agriculture or forestry experiments we do not expect different parts of a plot or field to vary systematically – one point is equivalent to any other point. This is not the case in typical agroforestry plots. Tree effects will not be uniform across a plot; for example, the soil properties under a hedgerow might be expected to differ from those under the crop between two hedgerows. The sampling used will depend on whether assessing this difference is required to meet the objectives.

Composite samples and bulking In most cases, the objective of a sampling programme is to obtain average values of certain soil characteristics for a given area. In this case, the soil from different sampling points is bulked, mixed and a subsample taken for analysis. If this is done, then the variation between repeated subsamples gives no information about variation in the field.

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In some cases it is necessary to take measurements on individual samples collected in the field. An example is the estimation of the precision of a plot average, usually the standard error of a mean of several samples in the plot. This information is useful for optimizing sampling schemes during preliminary studies but is of no use for the comparison of experimental treatments in different plots. Estimating precision is straightforward if the samples were collected in a random pattern. Systematic sampling does not allow precision to be estimated in a simple way, but estimates are available through the use of geostatistical methods to analyse the spatial patterns of soil properties in a plot (see Section 3.6).

Sampling uniform areas An area to be sampled (a field or experimental plot) that is uniform, with no known or predicted patterns of variation across it, is unusual in agroforestry research. Possible examples for uniform areas are the croponly control in an experiment, agricultural plots following a homogeneous fallow and perhaps very dense and homogeneous fallows themselves. The sampling scheme determines where samples will be located in the area. Two schemes for this situation are random and systematic. Random sampling ideally means that the coordinates for each sample location are selected using random numbers, but in practice the samples are usually collected on a zigzag path across the plot (James and Wells, 1990). In systematic sampling, the sample sites are selected in a systematic way, typically using a grid pattern. For example, a 20 m ¥ 20 m plot could be sampled by locating 16 points in a square 5 m ¥ 5 m grid, starting 2.5 m from the edge of the plot. Random sampling has the advantage that it is, in theory, unbiased, as no part of the plot is favoured. However, the practical application of random sampling can be biased as data collectors tend to avoid points they feel do not look ‘typical’. Systematic sampling has the advantage that it is unbiased except in the case of the grid points coinciding with some pattern in the field. The example above would clearly be inappropriate if trees were also planted on a 5 m ¥ 5 m grid. The practical application of systematic sampling is simpler than random sampling because the same sample positions can be used in every plot and the subjectivity found in practical random sampling is removed. Systematic sampling will nearly always give estimates of higher precision than random sampling because the sampling points are more evenly spread through the area of interest (Webster and Oliver, 1990).


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Sampling non-uniform areas Most areas to be sampled vary in a known or predictable way. The patterns of variation may be due to factors such as slope, soil depth or weed distribution. In agroforestry plots, pronounced variation of soil characteristics, litter and root distribution is often caused by the spatial arrangement of different trees and crops, and a common problem is how to obtain a sample that is representative for the whole plot in this situation. A useful approach to this problem is to identify the smallest representative unit (SRU), which is the smallest spatial element of which the plot (or system) is composed. For example, in a planted fallow with trees at 2 m ¥ 2 m spacing, the SRU would be a 1 m ¥ 1 m area with a tree in one corner (Fig. 3.3). In a system with trees planted in rows between crop fields with 5 m spacing between trees in the rows and 20 m between rows, the SRU would be a 2.5 m ¥ 10 m strip perpendicular to a tree row with a tree in one corner if the direction from the tree row is not considered important (SRU 1 in Fig. 3.4), or a 2.5 ¥ 20 m strip if there are directional effects of slope or prevailing wind that need to be taken into consideration (SRU 2 in Fig. 3.4). Even in a pure crop field the SRU concept may be useful, for example, when a field has been ridged, when a crop is still small and does not occupy the soil homogeneously or in most cases when root systems and related soil characteristics are studied. To be representative for the whole plot or system, a sample would have to be representative for one or preferably more SRUs in a plot. To obtain a representative sample of litter in an agroforestry plot, an efficient strategy can be to collect the whole litter in one or more SRUs per plot, then homogenize and subsample. For measuring coarse root biomass, the excavation of the soil of one or more SRUs per plot to a certain depth may

Fig. 3.3. Smallest representative unit (SRU) for sampling an area with trees planted at a spacing of 2 m ¥ 2 m.

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Fig. 3.4. Two options for the smallest representative unit (SRU) for sampling a crop field between rows of trees that are planted at 5 m ¥ 20 m spacing. The choice between the two SRUs depends on whether a significant influence of the direction from the tree rows on the variables under study is expected (SRU 2) or not expected (SRU 1).

be necessary. For soil or fine root samples, the individual sampling points would have to cover one or more SRUs in a plot in a representative grid or other pattern. In many cases, the precision of sampling can be increased, and additional information on spatial structures obtained, by stratification. This simply means dividing the plot into regions thought to be homogeneous, and sampling each of these strata. Within each stratum, either a random or a systematic sampling plan can be used. For example, in a pasture or crop field with scattered trees, the strata to be sampled separately could be: (i) the area influenced by the shade and litter of a certain tree species, and (ii) the area not directly influenced by the tree. In a hedgerow intercropping experiment, the plot could be subdivided into: (i) the area under the tree rows, and (ii) the area between the tree rows that is tilled, cropped and receives tree mulch. This cropped area could present gradients of nutrient availability caused by decreasing nutrient uptake by the trees and/or increasing nutrient uptake by the crops with increasing tree distance. In this situation, separate sampling of the soil under the trees and crops combined with systematic sampling within the cropped area (e.g. every 25 cm on a line crossing the area from one hedgerow to the other) ensures representative inclusion of soil from all tree distances. Stratified systematic sampling plans are usually the method of choice for root studies in agroforestry to allow for small-scale patterns in root mass and density around individual trees or tree rows.


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When a stratified plan is used but interest is in the whole plot, care must be taken to weight the observations from each stratum appropriately (Rao and Coe, 1991). The following sampling design for a 1 m ¥ 1 m tree plantation was used by Jama et al. (1998). The plot is divided into three strata according to how far each point is from the nearest tree. The strata of 0–0.28 m, 0.28–0.48 m and 0.48–0.7 m from a tree represent 25, 50 and 25%, respectively, of the area (Fig. 3.5). The choices now are: (i) sample each stratum, measure the soil property of interest for each and average the results using weights of 0.25, 0.5 and 0.25, respectively; or (ii) create a representative bulk sample from each stratum in the ratio 1:2:1. A sampling scheme with five instead of three tree distances for the same situation has been used by Mekonnen et al. (1999). Pronounced spatial patterns of soil fertility within a plot, which have to be taken into account when defining sampling schemes, result not only from the presence of different tree and crop species in a system, but also from standard agricultural practices such as fertilizer placement or certain tillage methods. Tree crops are commonly fertilized non-uniformly, for example, in bands along coffee rows (on sloping land on the upper side of the row) or in a circle around the stem of larger, more widely spaced trees (as is common for citrus, oil palm and other tree crops). The area where the fertilizer is applied is often kept free from ground vegetation to reduce nutrient uptake by weeds or cover crops. Over the years, this 1m 25% 50% 25% 1m Tree

Fig. 3.5. Schematic diagram illustrating the subdivision of a plot with trees planted at 1 m ¥ 1 m spacing into three sampling strata (reproduced with permission from Jama et al., 1998). Note that the 1 m2 square around the tree contains four smallest representative units and that for most research objectives it is better to collect one or two samples per distance per tree from several trees in a plot than many samples from a single tree.

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practice can lead to pronounced differences in chemical soil characteristics between fertilized (and weeded) and unfertilized (vegetation-covered) areas in a plot. For example, available soil phosphorus (Mehlich 3) at 0–10 cm soil depth in an Amazonian oil palm (Elaeis guineensis) plantation was 165 mg kg–1 at 1 m stem distance, but only 4 mg kg–1 at 2.5 m stem distance (Schroth et al., 2000a). In such situations, it is best to sample and analyse the fertilized and the unfertilized soil separately and to complement the analysis with a root distribution study to obtain an idea of the relative importance of these soils of differing fertility for plant nutrition. This example also illustrates how sequential samplings of the same plot could erroneously detect changes in soil fertility by collecting samples at slightly different positions if spatial fertility patterns are not taken into account. Tillage practices that lead to pronounced spatial fertility patterns within a field include ridging, a common system not only on sloping land, but also on level areas, for example in West Africa. With a specially designed plough or a hoe, a strip of soil is turned and moved sideways, where it covers a soil strip of similar width and forms the ridge. Topsoil and weeds are concentrated in the ridges, on which the crop is sown. Microerosion progressively takes fine materials from the ridges into the furrows during the growing season. If sample collection before the ridging is not possible, sampling halfway up the ridges has been recommended (James and Wells, 1990), but it is often better to collect and analyse two separate samples from the ridges and the furrows, especially if different crops are grown in these two positions.

Comparing parts of a plot In agroforestry experiments, plots are not only subdivided to avoid problems with sampling heterogeneous areas. In many cases, the analysis of the spatial patterns of soil properties that are caused by the different plant species is of considerable interest itself, as it may provide information on the different effects of trees and crops on the soil and so on tree–crop interactions. Examples would be systematic changes in soil nitrogen or soil water content with increasing tree distance in a crop field. Figure 3.6 shows a plot with a single row of trees down the centre with crops on either side (as would be appropriate for a shelterbelt). To measure the spatial variation of soil properties caused by the trees, samples have to be collected at different distances from the tree line. The principles are the same as for choosing quantitative levels of a treatment factor (see Section 3.2). If a simple response to distance is to be estimated, then it is better to use many samples at a few distances rather than few samples at each of many different distances. Typically, measurements will be taken in or close to the tree row (maximum tree effect), at a large distance (no tree effect) and


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Tree row Crop Sample transect

Fig. 3.6. Sampling scheme in transects in a plot with a single tree row and crops.

at one or two intermediate distances. Soil properties usually change most rapidly close to the trees, and it is thus advisable to collect samples at smaller distance intervals here and increase the intervals with increasing tree distance. Precision for comparing distances will be maximized if the samples fall in a number of transects, as in Fig. 3.6. In certain cases, measuring a representative value for a whole plot or system may not be the main objective of a sampling strategy. It may be more important, for example, to detect extreme values within a plot. When tree species that differ in their effects on the soil are mixed in a complex system, such as a forest garden, it would be expected that the effect of each tree is most pronounced close to the tree and decreases with increasing distance from the tree, where it also overlaps increasingly with the effects of neighbouring trees. In this situation, the most sensitive sampling strategy for the detection of specific tree effects on soil fertility is the collection of soil samples taken from very close to individual trees of the different tree species for comparison with controls taken at points at a maximum distance from all trees, which are assumed to have the smallest influence from trees possible given the configuration of the system. The resulting single-tree effects on soil fertility cannot be quantitatively scaled up to the whole plot unless extensive sampling is undertaken to characterize the gradients and geostatistical techniques are employed (see Section 3.6), but they do give valuable qualitative information concerning the way a certain species and its management might influence soil conditions, which is often of more immediate practical use than quantitative mean data per hectare (see Schroth et al., 2001b, for a review of agroforestry applications). In parklands and similar systems with naturally regenerated trees in irregular arrangements, care must be taken in the interpretation of observed fertility differences between tree and notree positions, as the trees may have established themselves on fertile spots, such as termite mounds or depressions receiving run-on water and eroded nutrients. The possibility of such initial fertility differences between positions may be excluded when the trees have been planted at a regular spacing, as in tree crop plantations.

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Sampling depth Soil sampling for agricultural purposes often includes only the topsoil, that is, the plough layer or a shallow surface horizon in no-till fields (James and Wells, 1990). In forestry, deeper sampling sometimes gives better correlations with plant growth or fertilizer response, for example 60 cm for phosphorus and 150 cm for potassium in different pine stands as reported by Comerford et al. (1984). For nitrate, sampling to 120 or even 180 cm depth has been recommended even for annual crops (Comerford et al., 1984; James and Wells, 1990). In agroforestry, sampling of the top 10 or 20 cm of soil may sometimes be adequate when treatments that are unlikely to affect the subsoil are being studied, such as the short-term effects of mulching, but it is inadequate for studies of competition and complementarity of nutrient use between different species. As a rule, the decision about sampling depth should be based on an at least qualitative analysis of the vertical root distribution of the relevant plant species (see Section 12.2). If this analysis reveals that one species (the crop) has relevant amounts of roots to 60 cm soil depth and the other species (the tree) to 100 cm depth, it is important to collect a separate sample from the 60–100 cm depth as the water and nutrients here may contribute to complementarity between the species (see Box 5.2, p. 100). If the tree penetrates a hardened subsoil horizon but the crop does not, the hardened horizon and underlying horizons should be included in the sampling, because increased access to the subsoil for the crop following root channels of the tree could be an important benefit of the association. If nutrient recycling from the subsoil is to be assessed, a preliminary evaluation of the nutrient distribution in the soil profile is useful. It is common practice to take fewer samples from depth than from surface layers to reduce the sampling effort. However, few samples will lead to imprecise estimates, unless the variability of the investigated variable also decreases in the subsoil.

Sampling time and frequency Certain soil characteristics change both during the year and between years, and sampling time has thus to be selected carefully in relation to the objectives of the study. Seasonal fluctuations have been reported for labile fractions of carbon, nitrogen and phosphorus as well as microbial soil characteristics in response to moisture changes, crop phenology and management (Conti et al., 1992; Mazzarino et al., 1993; Campo et al., 1998). Soil test readings for phosphorus and potassium probably decrease during the cropping season, whereas soil acidity increases in acid, but not in alkaline soils (James and Wells, 1990). In long-term studies, samples


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repeated in different years should be collected at the same time of the year relative to weather and crop development to be fully comparable (e.g. ‘2 weeks after crop emergence’ or ‘as soon as 50 mm cumulative rain has fallen after 1 September’). Short-term fluctuations of nutrient availability in response to management measures such as fertilization and mulching can also be studied by collecting and analysing the soil solution at appropriate intervals (see Section 7.2).

3.5 Analysing the Data (R. Coe) There are many resources that describe standard statistical procedures for analysing data from experiments. There are, however, few books that actually describe the process of analysing data from field trials, so this is discussed briefly here. The first step is to define objectives for analysis. The difficulty many scientists face with statistical analysis originates in part from the analysis objectives not being carefully thought through. Defining the objectives of analysis will involve: • •

•

identifying the exact comparisons to be made or relationships to be estimated; determining the exact data that are needed to make the comparisons (for example, are comparisons of yield needed each season or totalled over all seasons?); and designing the tables and graphs that will be used to present the results.

The objectives of analysis are determined by the objectives of the trial. However, the analysis objectives are distinct from trial objectives in that: • • •

•

•

the objectives of the trial may well have been stated in a rather vague way; new objectives will have been developed as the trial progresses, resulting from observations made; the objectives set out in the original protocol may well have other, unstated, objectives added if these can usefully be met with the data available; it might not be possible to meet all the original objectives of the trial, either because the trial design does not allow this, or because something unexpected has happened to prevent it; and the objectives of the analysis will evolve as the analysis proceeds.

The second step requires preparation of the data. Almost always there is considerable work to do to turn raw field records into data files suitable for statistical analysis. Conversions and calibrations have to be completed. New variables have to be constructed. Data have to be summarized to the

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appropriate space and time scale, and so on. The ease with which this is done depends largely on how the original records were organized. Statistical Services Centre (2000) is a good guide to optimizing this. The third step is exploratory analysis of the data that aims at finding summary tables and graphs that meet the objectives. Well-constructed tables and graphs will make the patterns and relationships in the data clear and should make tentative conclusions obvious. The other aim of exploratory analysis is to reveal any unexpected observations or patterns in the data that might impact either on the conclusions or the way the analysis proceeds. Next comes the confirmatory analysis. This is where formal statistical techniques are relevant and is the step widely described in books and taught in courses. The formal analysis aims to: •

• •

confirm that the patterns and relationships noted in the exploratory analysis are consistent with real or repeatable effects, and are unlikely to be due to noise; estimate the precision of important quantities emerging from the analysis; and increase the precision of these estimates by finding parsimonious statistical models which describe the data, do not contain any unimportant terms and manage to explain much of the variation.

The final step involves interpretation and reporting. Iteration between the various steps of analysis is nearly always needed. Data from agroforestry-fertility field experiments are not inherently different from data from other trials, so these general methods usually apply. However, there are some characteristics of data from agroforestry trials which can make it confusing to apply general methods.

Multiple outputs In system trials aimed at evaluating the productivity of alternative systems there will often be multiple products (tree and crop products, for example). The options for analysing these are to look at them separately, to investigate the relationship between them and to combine them into a single index of production. The range of methods developed for analysis of intercropping trials are appropriate (Federer, 1993, 1998).

Measurements repeated in time The time scale of agroforestry experiments means that measurements will almost inevitably be taken repeatedly on the same variable at several times.


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These may be within one season, for example to look at nitrogen dynamics during a cropping phase, or across seasons to look at long-term development of systems or response to different weather. The statistical problem associated with repeated measures arises from the fact that observations on the same units at different times cannot be expected to be independent, with correlations between observations likely to be larger the smaller the time interval between them. The most common statistical procedures, such as those based on analysis of variance (ANOVA) and regression, require independence of observations. A large group of techniques under the general heading of repeated measures analysis has been developed to handle this problem. These techniques range from simple and approximate adjustments of standard analyses to statistical models that describe the correlations in detail. They are described in many textbooks and appear in standard software. However they can be difficult to apply, particularly if the times of measurement are irregular or different for different treatments. There is one approach which is simple to apply and produces analyses which are well tuned to meeting objectives. The idea is to turn the repeated measurements on each experimental unit into a single value for each unit, that value being chosen to represent a quantity required by the analysis objectives. The steps are as follows. 1. Explore the data (usually by plotting the results for each treatment against time) in order to identify a pattern or effect that is of interest and needs formal statistical analysis. 2. Choose a number that describes the effect of interest. Examples include: • the total biomass produced over 5 years; • the difference in biomass produced in a wet season and a dry season; • the change in soil nitrogen between crop planting and leaf stage 6 minus crop nitrogen uptake, representing nitrogen losses in the early part of a season; • the time until soil carbon reaches some threshold level, based on interpolation of annual measurements; • the rate parameter k of a decomposition curve fitted to biomass decomposition data. These examples show that the summary can be something as simple as a total or difference, or as complex as the parameters of a non-linear curve fitted to trends. The important point is that the number describes the phenomenon of interest. 3. Calculate the number for each experimental unit or plot of the original design and analyse for treatment differences using the ANOVA dictated by the design.

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4. Repeat the procedure for as many different summary numbers as are needed to meet all the objectives.

Measurements repeated in space Repeated measurements also occur in space, as different parts of a plot are measured with the purpose of comparing, but they do not represent separately randomized treatments. In many soils experiments, measurements are taken at different depths. Agroforestry experiments often have the added complication that different positions within a plot are not equivalent, for example different distances from a line of trees. The statistical problem associated with these repeated measures in space is identical to that posed by repeated measures in time – the observations from different parts of a plot cannot be expected to be independent. The approaches for dealing with the problem are also similar, though the statistical modelling of correlation structures in space can be rather harder than the equivalent in time. The simple approach based on summary quantities is also applicable to repeated measurements in space. For example, suppose tree and crop root length densities are measured at five different distances from a tree line and at eight different depths within each plot of an experiment. Depending on the exact objectives, examples of suitable summary values to be calculated for each plot and subjected to formal analysis might be: 1. Proportion of tree roots below the crop root layer; 2. Total tree roots per metre of tree line; 3. Root length density of crops in the zone 0–50 cm deep and less than 3 m horizontally from the tree line; 4. Parameters A, k and c in the model A exp(–k(h2+cv2)0.5) that describes the change in tree root length density with horizontal (h) and vertical (v) distance from the base of the trees. It is likely that the summary chosen cannot be calculated in the same way for all treatments. Suppose the examples above come from a trial that compared four different tree species grown in a crop, together with a croponly treatment with no trees, a total of five treatments. Summary (1) is simply not defined for the crop-only treatment. When analysing this variable there are only four treatments to consider. Summary (2) makes sense for the crop-only treatment but has the fixed value of zero. Since there is no uncertainty in this treatment mean, it is not correct to include these zeros in the analysis along with other values. When comparing a mean from a tree plot with that of the control we are comparing an uncertain non-zero mean with a known value of zero. For summary (3) we may well want to compare the value from tree plots with that from crop-


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only plots. The value for the crop-only plot will be calculated from all measurements 0–50 cm deep in the plot, as different locations horizontally are all equivalent. The fact that the summary is calculated in different ways in plots of different treatments does not affect the formal analysis, apart perhaps from inducing some non-constant variance, which can be allowed for by suitable weighting. Software for experimental design No software will design experiments for you. However, there are several components of experimental design for which software can help. Software to help estimate the number of replicates or sample sizes needed is reviewed by Thomas and Krebs (1997). Their online paper also gives links to providers of commercial and free software. Design of efficient incomplete block designs, neighbour designs and other advanced designs that allow for some site heterogeneity is assisted by the CycDesign software from the Commonwealth Scientific and Industrial Research Organisation (CSIRO), Australia (www.ffp.csiro.au/ tigr/software/cycdesign). Similar facilities are available with the Genstat programme (www.nag.co.uk). There are many software products available aimed at helping to design industrial experiments. These are mainly concerned with selection of efficient treatment combinations from very large multifactor sets and have little application in agroforestry research. Software for statistical analysis There are hundreds of products to help in statistical analysis of data from experiments. The number around is now so large that there seems to be no systematic and reasonably complete list or directory to them. Internet sites of many statistical organizations contain links to software sites. One extensive listing that may be useful is found at www.stats.gla.ac. uk/cti/links_stats/software.html#packages. Many of the products listed here are accompanied by independent reviews. In addition to statistics packages there are facilities for certain statistical manipulations in many other types of software – spreadsheets, databases, even graphics software. With such a large choice it is difficult to make recommendations, though some general comments may be useful. First, it is sensible to use a dedicated statistics software package for statistical calculations. An excellent article explaining the limitations of the spreadsheet Excel for statistics, for example, is found at www.rdg.ac. uk/ssc/dfid/booklets/topxfs.html. Similar points could be made about trying to do anything other than the simplest statistics with, say, database or graphics systems.

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Secondly, there is no statistical software that meets all the requirements of any analyst. For example, S-Plus (www.splus.mathsoft.com) has many of the more recent developments in statistical methodology incorporated, but has a steep learning curve and is not easy for beginners to start using. Minitab (www.minitab.com), on the other hand, is ideally suited to beginners but may well not be able to analyse some of the more complex designs found in agroforestry research. Generally you get what you pay for. It is unlikely that you will find free or shareware products that allow you to complete effective analyses. However, cost is not a guarantee of quality or appropriateness for analysis of experiments, and software which is sold as a general statistical analysis system may not be particularly suited to analysis of experiments as opposed to surveys, for example, SPSS (www.spss.com). Possibly the only freely available statistical software that does make a suitable introductory package for experimenters is INSTAT (www.rdg.ac.uk/ssc/instat/instat.html), which comes with extensive high quality documentation and examples. SAS (www.sas.com) is often considered as the standard against which other statistical software is judged. It certainly has very extensive facilities and is widely used. Because of this, analyses using SAS are described in many books. However, there are some disadvantages. It can be difficult to carry out some non-standard analyses or manipulations that are conceptually straightforward. More importantly, SAS has developed a system for maintaining, integrating and delivering information from a wide range of sources. The applications development and database facilities have become very advanced, but will probably not be exploited by the agroforestry researcher, who ends up having to pay for software that will not be used. The decision on whether or not to invest in SAS will generally be an organizational rather than an individual one. Many less-common analyses will require special software. Software for spatial statistics is reviewed in Section 3.6. Other examples of possible relevance are MIM (www.hypergraph.dk) used to unravel complex multivariate dependencies, or PC-ORD (www.ptinet.net/~mjm) for analysis of species composition data. Given the great choice in software available and the range of tasks the analyst of agroforestry trials has to carry out, the only strategy that can be recommended is to acquire one of the well known and general statistical analysis systems and become proficient at using it, but recognize that additional software may be needed for particular jobs. The choice of which main package to use will often depend on who else is using it and what other tasks, beyond analysis of agroforestry experiments, it will be used for. An example of software which allows straightforward analysis to be completed easily, leads naturally to more complex analyses and overall maximizes the chance of doing a sound analysis is GENSTAT (www.nag.co.uk).


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3.6 Spatial Structure and Its Analysis (B. Huwe) There are several aspects to problems associated with spatial heterogeneity or variability in agricultural and agroforestry systems. On the one hand, field variability is regarded as a major obstacle in identifying parameter impacts or relationships between system components. High variability obliges the experimenter to increase the number of replicates in the experimental design or to subdivide the study area into several, separately sampled strata, which increases the number of samples and the complexity of the data analysis (see Section 3.4). On the other hand, the spatial variabilities of soil, vegetation and other properties are important system characteristics and thus of interest themselves. For example, the detection of spatial correlation between the distribution of plant species or their root systems and soil properties may be a first step in the analysis of plant–soil interactions (Adderley et al., 1997; Mekonnen et al., 1999). Further, in many applications extreme values can be more important than mean values, such as leaching in macropore systems, influenced by tree roots or macrofauna (Beven, 1991), or denitrification in ‘hot spots’ such as local clay enrichments or soil aggregates with reduced aeration (Clemens et al., 1999). Spatial patterns and probability distributions are essential in studies where uncertainty analyses are involved, such as the risk of yield failure in drought years as influenced by variable soil texture or soil depth in a landscape, groundwater pollution with pesticides or nitrate (Soutter and Pannatier, 1996; Wade et al., 1996), or N2O emissions into the atmosphere (Velthof et al., 1996; Clemens et al., 1999). This section focuses mainly on geostatistical concepts, which means spatial analyses and predictions, and their potential applications in agroforestry research. Limitations of geostatistical methods are pointed out, and some alternatives are mentioned. For procedures that allow the simultaneous treatment of space–time systems, see Myers (1992) and Panesar (1998). A short and incomplete list of available software is given at the end of this section.

Potential use of spatial statistics Geostatistics provides useful tools for spatial analyses, creation of optimized maps and simulation of spatial processes. For example, geostatistics can be used to obtain estimates of the total nitrogen accumulation in the soil under a legume fallow, or total phosphorus availability in the soil of an agroforestry plot with complex spatial patterns of fertilizer application and nutrient uptake by plants. When creating maps of chemical or physical properties of a field or landscape, information on the similarity of values from neighbouring samples and increasing dissimilarity of values with

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increasing distance between the samples, the so-called autocorrelation structure for the area, can be used to calculate weighting factors which minimize the estimation error of the variables under study (Goovaerts, 1998). Geostatistical methods can also be used for the generation of optimized sampling schemes according to the requirements of the experiment, financial constraints or predefined local accuracy (Webster and Burgess, 1984), thereby reducing the experimental effort. Further, it is possible to identify optimal pixel sizes for process models (Wade et al., 1996) or to generate parameter fields for the analysis of the transport of nutrients or pollutants in structured soils (Piehler and Huwe, 2000). Lopez and Arrue (1995) used geostatistics to optimize an incomplete block design for tillage experiments which proved to be considerably more efficient than the corresponding complete block design. Although geostatistical methods have been widely used in the field of soil, agricultural and hydrological sciences at different scales (Oliver, 1992; Schiffer, 1992; Webster and Boag, 1992; Huwe, 1993; Hoosbeck and Bouma, 1998), not much work on spatial analysis has been reported so far in the field of agroforestry. However, potential benefits for agroforestry studies are demonstrated by applications in forest and agricultural ecosystems (Sylla et al., 1996; Bragato and Primavera, 1998; Gorres et al., 1998), orchards (Gottwald et al., 1995), shrub–steppe ecosystems (Smith et al., 1994; Halvorson et al., 1995) and in soil surveys (Di et al., 1989). The scale of application ranges from the microscale of soil pores (Grevers and Jong, 1994) through agricultural fields (Bragato and Primavera, 1998) and landscapes (Wade et al., 1996) to regions of several thousand square kilometres (van Meirvenne et al., 1996). Processes involved in the studies comprise tree growth (Meredieu et al., 1996), crop yield (Wopereis et al., 1996), water transport in soils (Mulla, 1988), evaporation (Lascano and Hatfield, 1992), carbon and nitrogen mineralization (Smith et al., 1994), nitrous oxide fluxes (Velthof et al., 1996) and pesticide transport to the groundwater (Soutter and Pannatier, 1996). As water and matter fluxes play an increasing role as indicators for sustainability and environmental compatibility, the aforementioned aspects of heterogeneity will be of increasing importance in future agroforestry studies. The epidemiology of plant diseases and even the spatial distribution of leaves and fruits in trees have also been studied by means of geostatistical methods (Chellerni et al., 1988; Monestiez et al., 1990; Orum et al., 1999).

General principles of geostatistics Spatial variability, or spatial patterns, can conceptually be divided into components which are caused by either deterministic or stochastic processes. Apparently stochastic behaviour of a system may be induced by


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so-called hidden variables (i.e. variables that were not measured), or simply by the scale of measurements. For example, to predict nitrate leaching in the soil, we may use an average root length density and average hydraulic conductivity per soil layer as determined from a number of field measurements as input variables into a deterministic model. However, even in the best of cases the output of this model will not be correct for every single spot in the area, because not every spot has average properties with respect to root length density or hydraulic conductivity and, of course, we can never know the exact root length density and hydraulic conductivity at every spot in the soil. Thus, whereas part of the factors that influence nitrate fluxes are ‘known’ (i.e. have been described through measured, average values), another part will always remain unknown and will introduce unexplained (apparently stochastic) variability in the behaviour of the system. As mentioned before, this unexplained variability can be so large in certain cases that the model result will be completely wrong, simply because the behaviour of the system is not determined by average but rather by extreme values. The identification, analysis and description of the deterministic system components belong to the field of deterministic, process-based modelling, whereas the analysis and modelling of stochastic patterns is done within the framework of the theory of stochastic processes, including geostatistical theory. Classical statistical methods which are generally used in agroforestry largely ignore spatial structures. Most tests assume independence of sampling points and, in addition, normally distributed variables or residuals. By ignoring spatial autocorrelation structures, an error is introduced into the analysis. The fundamental backbone of geostatistics is the concept of regionalized variables, which is based on the work of engineers, mathematicians, physicists and biometricians in the early 1940s (Mathes and Ries, 1995). The theory in its present form was developed mainly by Matheron (1963). For original literature and more complete descriptions of geostatistical theory and applications see Journel and Huijbregts (1978), Journel (1989), Bárdossy (1992) and Webster and Oliver (2001). The main idea behind geostatistics is the common observation in the field that values of variables often resemble each other more as the distance between sampling points decreases. With increasing distance, the spatial influence of neighbouring samples becomes smaller, and above a certain limit, the so-called range, variables are independent in a statistical sense. Measurements are treated as realization of an ergodic spatial stochastic process, which means that the characteristics of the stochastic pattern or process can be derived from this single spatial data set. For example, a single measurement of soil phosphorus in several sampling positions in a plot is enough to describe the stochastic pattern of phosphorus distribution in the plot. Another assumption of geostatistical theory is intrinsic (or weak)

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stationarity, which means that for any two sampling points in the area, the difference in the measured value (e.g. soil phosphorus content) depends only on the distance between the two points (the lag), independently of where in the plot the two samples were collected. This assumption is often problematic, especially in areas with pronounced soil gradients or abrupt changes in soil properties, as will be discussed further below. Procedure of geostatistical analysis A basic component of geostatistical analysis is the semivariogram (Fig. 3.7). A semivariogram is a mathematical function that describes how two sampling points become more different with increasing spatial distance between them, until a certain distance when the points become independent from each other. The semivariogram describes the autocorrelation structure of the measured variable in the respective area. The mathematical formula of the semivariogram is given further below. Geostatistical studies involve the following steps (Kitanidis, 1997): • • •

design of the sampling grid; data collection; analysis of spatial structure (determination of empirical semivariograms); range 10

Semivariance

8

6 sill 4

2 nugget variance 0 0

2

4

6

8

10

12

Lag

Fig. 3.7. Schematic semivariogram with a nugget variance of 2, a sill of 7 and a range of 7.


• • •

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selection of an appropriate semivariogram model; interpolating the grid by best, linear, unbiased estimations (BLUES); and interactive optimization of the semivariogram model using crossvalidation techniques.

The design of the sampling grid (dimension, grid size, regular or irregular spacing) depends on problem-specific accuracy requirements and the spatial structure of the sampling area. One-dimensional grids (transects) are used for linear structures, such as gradients between linear tree plantings and adjacent fields. Two- and three-dimensional grids are used for maps, for example of nutrient distribution in a plot, or for the determination of parameter fields for process simulations, such as the transport of nutrients and pollutants in soil. Webster (1985) found that for isotropic semivariograms, i.e. in areas where soil properties are expected to change according to the same pattern in all directions, the best twodimensional sampling scheme is a regular, equilateral triangular grid. For the estimation of mean values of square blocks, the best position of sampling points is in the centre of the blocks (Webster and Burgess, 1984). However, in practical studies it is often not possible to assure a regular grid. Further, in many cases it is advantageous to use a nested grid in order to determine the semivariogram for short sampling distances (Selles et al., 1999) and to focus the accuracy of the calculated map on a predefined area of special interest (e.g. a sampling plot within a larger section of the landscape). A qualitative explanation of the semivariogram has been given above. The mathematical formula of the semivariogram g( → h ) is given by r

()

g h =

1

r 2N h

()

r N ÊË hˆ¯

Â ( z(xi) – z(x i + h)) r

r

r

2

(3.2)

i=1

where z(→ x ) denotes the measurement at location → x , and N is the number of pairs with a distance of → h . The arrow indicates vectors. In simple words, the semivariogram gives for every distance h between two data points a mean squared difference, obtained by averaging all pairs of data points with this distance in the data set. For a small h (samples collected close to each other), this difference will be relatively small, and for a larger h it will be greater. The use of vectors in the formula indicates that the semivariogram may not be the same in all directions, i.e. it may be anisotropic. As illustrated in Fig. 3.7, typical semivariograms are characterized by: (i) a short-range variability, the nugget variance, which is determined by the closest sampling distance in the grid and is therefore to a certain extent an artefact; (ii) an asymptote equal to the background variance of the

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variable, i.e. the variance between sampling points that are completely independent of each other at the scale of the study; and (iii) a maximum lag value that characterizes the range of spatial influence and is therefore called range. Points that are farther apart than this range are independent at the scale of the study. The sill is the difference between the background variance and the nugget variance. For irregular grids and anisotropic conditions, the differences are grouped into several distance and angle classes. The experimental semivariogram is the basis for the selection of a theoretical model, which is then used for the analysis of spatial structures of the variable under study. Theoretical semivariogram models are mathematical functions that resemble the empirical semivariogram as closely as possible. Not all functions that would fit the data can be used as semivariogram models. If the model is to be used for spatial estimates or spatial simulations, it must be positively defined, a mathematical requirement that will not be discussed here (see Journel and Huijbregts, 1978). Unfortunately, it is not trivial to prove whether a given function fulfils this requirement or not. Custom models that are known to fulfil this criterion are the linear, spherical, exponential, logarithmic and Gaussian models. Other positively defined models are the quadratic, rational quadratic, power and wave models (Alfaro, 1980; Cressie, 1991; Pannatier, 1996; Kitanidis, 1997). Models for anisotropic conditions are also available. The field of variography is still in development. A first estimate of semivariogram model parameters can be obtained by a least squares approach using the empirical semivariogram data. In a subsequent procedure, a number of different kriging techniques (see below) can be used to optimize the semivariogram and yield the bestpossible estimated data. A criterion for the quality of the estimation is obtained by jack-knifing, a cross-validation procedure where each measured point is estimated from the neighbouring points and the estimated values are compared with the true values. The correct optimization algorithm for the semivariogram thus includes the whole estimation procedure, including the krige algorithm itself (Kitanidis, 1997), and not only the fitting of measured and estimated semivariogram data.

Point kriging Kriging is a linear unbiased estimator for the variable under study. It is calculated as the weighted mean of the values of all sampling points or of the points in a user-defined local neighbourhood (which often ascertains local stationarity and thus avoids trend corrections, see below): ˆz =

Â l i ◊ zi i

(3.3)


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where zi denotes the measured values, ˆz the estimate at a user-defined location and li the weighting factors calculated from the krige system. Kriging is also BLUES – the best, linear, unbiased estimator. Kriging, by definition, minimizes the estimation variance, which is the variance between the measured and the estimated data points. For each estimation on a location between the grid nodes, kriging involves the solution of a linear equation system, the krige system, that yields the weighting factors li for the linear estimation procedure and a so-called Lagrange multiplier m, which results from the consideration of the constraint l i = 1 . With the ls

Â i

and m it is possible to calculate the estimation variance for each estimated value and thus to determine confidence intervals. If the stochastic assumptions of the krige procedure are fulfilled, we may thus obtain not only maps with estimated (rather than mean) values, but in more or less the same step also maps that show the reliability of the estimated data, a source of information that is not provided by any other interpolation procedure.

Special kriging techniques Point kriging, as described above, is the simplest kriging technique and for many applications in agricultural or environmental sciences it is sufficient. For example, maps of soil texture or pH can be generated with this technique. There are also several advanced kriging techniques for special data structures, quality aspects and project goals that are briefly introduced below.

Trends A serious violation of the intrinsic model assumption of weak stationarity is given by a spatially non-constant expectation of the random variable under study, i.e. a trend. Trends may be caused by gradients in soil conditions in sloping areas, gradually changing geological or climatic site conditions, or land-use history. Common techniques to avoid the associated errors are the moving neighbourhood algorithm and the universal kriging approach. While the moving neighbourhood technique assumes local stationarity and thus restricts the grid points used for estimation to this neighbourhood, universal kriging assumes a predefined functional behaviour of the trend and eliminates this trend globally in an iterative algorithm (De Marsily, 1986; Deutsch and Journel, 1992). These techniques are available in some of the software packages listed at the end of this section.

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Block kriging Sometimes it may be desirable to estimate not point values but values that represent the average of the variable in a defined volume around the estimation point. Examples include the estimation of nutrient stocks or plant-available water in a given soil volume. The average variable is obtained by integrating over a volume V: z=

Ú z( x )dV r

(3.4)

V

The estimation process is analogous to point kriging and differs mainly in the averaging of the semivariances in the system of linear equations. As a consequence of averaging, the estimation variances are typically much smaller than those obtained by point kriging (De Marsily, 1986; Deutsch and Journel, 1992). Gorres et al. (1998) used block kriging for the generation of maps of carbon mineralization, bulk densities and nematode densities in a forest and agricultural field.

Co-kriging In most field studies, a number of parameters are measured at each sampling point rather than a single value. Many of these parameters may be intercorrelated as well as autocorrelated. Thus, taking into account the complete correlation structure should yield better estimation results compared to separate kriging of every single variable. Co-kriging was used by Halvorson et al. (1995) to analyse the spatial relations between resource islands in the soil and different plant species in a shrub–steppe ecosystem, and similar applications in savannas and parkland systems could be useful. As in simple kriging, co-kriging minimizes the estimation variance and provides its value for each location. Extensions to more than two correlated variables are possible (De Marsily, 1986).

Indicator kriging This is a technique that may become particularly useful in environmental decision problems, such as when an environmental measure depends on the percentage of values above or below a given threshold in the model area (Akin and Siemes, 1988; Mulla and McBratney, 1999). Indicator kriging requires the transformation of the variables into indicator variables (values above a defined threshold are assigned a value of 1, values below the threshold are assigned a value of 0). Variography and kriging are conducted as described above. Kriging is carried out as simple point


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kriging. Spatial averaging of the kriged indicator values yield the proportion of values of the original data set that are below the threshold. Halvorson et al. (1995) used this technique in the aforementioned study to determine the probability that certain combinations of variables occurred at unsampled locations.

Geostatistical simulation Kriging is an interpolation algorithm that is optimal in the aforementioned sense. However, it generates more or less smooth maps that are in some ways unrealistic when compared with the real situation in the field. Unfortunately, this not only is of interest from an aesthetic point of view but may even yield incorrect results. This is always the case when extreme values are of importance. For example, transport of nitrate or pesticides in soil is often governed by preferential pathways, caused by macropore systems or spatial heterogeneity of soil hydraulic properties. Using simple similarity concepts together with geostatistical simulation, it is possible to generate heterogeneous parameter fields. Effects of this simulated heterogeneity for transport processes may then be calculated with numerical transport models (Piehler and Huwe, 2000). For example, a critical parameter of denitrification in soils is the soil water content. As kriging is a smoothing estimator, the percentage of water content events above a critical value, and consequently denitrification and the production of nitrous oxides, may be underestimated. Conditional geostatistical simulation provides a measure to generate more realistic maps that are compatible with the determined semivariogram. Furthermore, in the grid nodes the simulated values are identical to the measured values. Several algorithms for spatial simulation are available (e.g. the random coin method by Alfaro, 1980). A comprehensive overview with algorithms and FORTRAN-codes is given in Deutsch and Journel (1992).

Alternative methods and tools Geostatistical methods have some serious drawbacks. Although normal distribution of the random variable is not explicitly required, it is recommended by some authors and may facilitate the interpretation of the maps of estimation variances. The applicability of geostatistics is restricted to situations where ergodicity may be assumed and the intrinsic hypothesis is valid. Thus, trends and structural discontinuities render the geostatistical analysis more difficult. Such situations may be caused by short-range, sharp gradients in a structured agroforestry system as well as by marked changes across a landscape (e.g. the boundary between

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grassland and forest), and geostatistics may then not be the appropriate methodology. Recent developments in mathematics and statistics provide algorithms and tools that also have potential for spatial statistics, although up to now they have not been studied and tested in full detail. Most of the methods have in common that they are robust and do not prescribe probability density functions like the bell-shaped Gaussian distribution. See Mathes and Ries (1995) for local gradient analysis, Batchelor (1998), Levine et al. (1996) and Liescheid et al. (1998) for artificial neural networks, and Woldt et al. (1992) for a geostatistical application of fuzzy set theory.

Geostatistical software Some information is given below about software available in the public domain (PD) and commercially (®). •

•

•

•

• • • •

GEO-EASPD and GEOPACKPD are DOS-programs that provide basic functionality of variogram analysis, kriging and visualization (‘rough and dirty’). The DOS-environment is not very user friendly and does not fit into modern Windows systems (www.hydroweb.com). SURFER® (Golden Software, 1999) is a contouring and surface mapping package that provides variogram analysis, variogram fitting and kriging, including block kriging and universal kriging (www.goldensoftware.com). GSLIBPD (Deutsch and Journel, 1992) is a most powerful collection of FORTRAN-programs that covers almost every aspect of geostatistics. The handbook is a very good overview of geostatistical theory. The user must be familiar with FORTRAN and have access to a FORTRAN compiler. SYSTAT® is a commercial statistical package that now includes geostatistical routines based on the GSLIB package and CARTAlgorithms (www.spss.com/software/ science/SYSTAT). GS+® is a commercial geostatistical software package with a user friendly interface (www.gammadesign.com). DATA-ENGINE® provides algorithms for fuzzy control, artificial neural networks and combinations of both (www.mitgmbh.de). Stuttgart Neural Network Simulator (SNNSPD) is a very powerful artificial neural network (www-ra.informatik.uni-tuebingen.de/SNNS). VARIOWIN (Pannatier, 1996) is an easy to use software package for variogram analysis and visualization.

Chapter 4 Soil Organic Matter G. SCHROTH,1 B. VANLAUWE2

AND

J. LEHMANN3

1Biological

Dynamics of Forest Fragments Project, National Institute for Research in the Amazon (INPA), CP 478, 60911–970 Manaus, AM, Brazil; 2Tropical Soil Biology and Fertility Programme, PO Box 30592, Nairobi, Kenya; 3College of Agriculture and Life Sciences, Department of Crop and Soil Sciences, Cornell University, 909 Bradfield Hall, Ithaca, NY 14853, USA

4.1 Synopsis Soils contain a variety of organic materials, ranging from living roots, fauna and microbes, through dead tissues in various stages of decomposition, to relatively stable, dark-coloured transformation products with no discernible anatomical structures, the so-called humus (Jenkinson, 1988b). Following Jenkinson (1988b), the term soil organic matter includes all organic substances in soil, both living and dead, although the living organisms (the soil biomass) usually contribute less than 5% to the total soil organic matter. Certain terminologies do not include the biomass in soil organic matter, and some authors use the terms humus and soil organic matter synonymously, so care is necessary to avoid misunderstanding. In practical terms, analyses of soil organic matter are usually carried out on air-dried soil that was passed through a 2 mm sieve and will include all living and dead organic materials in the soil that are not removed by this procedure (Anderson and Ingram, 1993). In studies relating soil organic matter contents or fractions to soil aggregation, particles >2 mm may also be included in the analysis. The organic materials that lie on the soil surface, including naturally fallen litter, crop residues, mulch materials and their decomposition products, are normally sampled and analysed separately from the soil (see Chapter 6). Because decomposing roots have a function similar to that of above-ground litter in the replenishment of soil organic matter, they are sometimes called below-ground (or root) litter. This should not be confused with the term soil litter, which has been used for a density/particle size fraction of soil © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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organic matter (see Section 4.3). A detailed discussion of the chemistry, formation and transformations of soil organic matter is provided by Stevenson and Cole (1999). The organic carbon contents of terrestrial soils vary between 100 µm, 150 mm, 53 mm) also confirmed the value of this fraction as an indicator of high nitrogen availability (Vanlauwe et al., 1998b). In a set of alley cropping trials in the West African moist savanna, the relative change in nitrogen in particulate organic matter was about twice the relative change in total soil nitrogen content, and the proportion of the total soil nitrogen in particulate organic matter increased significantly with annual nitrogen inputs in crop residues and tree prunings. This also suggests that nitrogen incorporated in this fraction is relatively labile and a more sensitive indicator of management effects than total soil nitrogen (Vanlauwe et al., 1999). Despite these promising results, it should be kept in mind that only part of the light fraction or particulate organic matter consists of readily mineralizable materials, another part consists of recalcitrant materials, whose relative contribution increases with progressive decomposition of added plant materials in the soil (Palm et al., 2001; see Section 4.3). Also,

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in many cases the light or particulate organic matter fraction is too small to account for all the readily mineralizable nitrogen in the soil (e.g. 2–3% of total soil nitrogen in the study by Barrios et al., 1997) and it does not include labile pools such as microbial biomass which is mainly associated with clay-sized fractions (Palm et al., 2001).

Chemical fractionation methods These difficulties could possibly be overcome by combining physical fractionation methods with chemical indices of nitrogen availability. An approach to quantifying a labile fraction of soil organic matter through selective oxidation with permanganate has been discussed in Section 4.4. Acid permanganate has also been used to extract ammonium-N from soil through selective oxidation of soil organic matter (Stanford and Smith, 1978). The extracted amount was found to be closely correlated with potentially mineralizable nitrogen and thought to represent a fraction of soil nitrogen that is readily susceptible to biological mineralization. Gianello and Bremner (1986) found a close correlation between ammonium-N extracted by boiling the soil in 2 M KCl for 4 h and potentially mineralizable nitrogen for 33 Brazilian soils of widely differing clay and organic matter contents. Egoumenides (1990) separated the total nitrogen in a large number of tropical soils by acid hydrolysis (6 N HCl for 16 h) into a non-hydrolysable fraction (mainly phenol and quinoneamino acid complexes), a hydrolysable-distillable fraction (mainly hexosamines, amides and certain amino acids) and a hydrolysable-nondistillable fraction (mainly amino acids of microbial origin). Greenhouse and field studies showed that nitrogen is supplied to plants principally from the hydrolysable-non-distillable fraction, which decreases under cropping and is sensitive to management measures such as crop rotations and fallowing. The hydrolysable-distillable fraction, in contrast, is mainly influenced by soil conditions, such as clay content. Numerous other chemical methods for the assessment of labile organic nitrogen have been described (Keeney, 1982).

Microbial biomass nitrogen The microbial biomass is another labile pool of soil organic matter and organic nitrogen as discussed in Chapter 4. However, nitrogen in microbial biomass is only sometimes correlated with nitrogen mineralization in the soil. This is not surprising, as the microbial nitrogen pool is too small to account for the amount of nitrogen mineralized from soil (Palm et al., 2001), and mineralization of nitrogen from other organic materials results

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from microbial activity rather than its biomass (Jenkinson, 1988a; Groot and Houba, 1995). The high temporal variability of microbial biomass nitrogen requires attention when defining sampling plans, and repeated measurements during a cropping season may often be necessary to obtain a representative picture (Haggar et al., 1993; Mazzarino et al., 1993). Care is also needed in the selection of methods when attempting to relate microbial biomass nitrogen to nitrogen mineralization across different soils. In a comparison of different grassland soils, Hassink (1995) found that nitrogen mineralization in the soil increased with nitrogen in the total microbial biomass as measured with the fumigation–incubation method, but that the amount of nitrogen mineralized per unit of microbial biomass nitrogen was higher for sandy than for fine-textured soils. In contrast, the relationship between nitrogen mineralization and nitrogen in the ‘active’ microbial biomass as measured by substrate-induced respiration was independent of soil texture. This effect of soil texture, which was confirmed for other soils by Franzluebbers et al. (1996), was explained by differences in physical protection of soil organic matter in coarse- and finetextured soils, and the second method was recommended for comparisons across different soil types. Detailed measurement procedures for microbial nitrogen are given in Anderson and Ingram (1993) and Voroney et al. (1993).

5.3 Methods for Soil Phosphorus ( J. Lehmann) Phosphorus availability is a limiting factor for plant production in many agricultural soils (Fairhurst et al., 1999). This is especially true in the highly weathered soils of the humid tropics. At the same time, the global availability of phosphorus fertilizers is limited and known reserves may be exhausted in about 100 years with the current growth of phosphorus usage (Stevenson and Cole, 1999). In regions of the world without a history of use of phosphorus fertilizers, phosphorus deficiency is very common (Wild, 1988). A large portion of applied fertilizer phosphorus may be fixed to iron and aluminium oxides and is then not available for plant uptake. These facts make sound phosphorus management an imperative, especially in situations where funds for fertilizer purchases are limited, as in tropical smallholder agriculture. Agroforestry techniques can help to overcome some of these constraints (Buresh, 1999). However, because of generally low phosphorus concentrations in mulch materials (see Chapter 6), low atmospheric inputs (see Chapter 9) and low release by mineral weathering, adequate applications of phosphorus fertilizer are necessary in permanent agriculture to ensure economic and ecological sustainability (Buresh et al., 1997; Newman, 1997). One-time replenishment of the soil


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phosphorus capital, e.g. with phosphate rock, has been proposed as an option for soil fertility management in impoverished soils (von Uexküll, 1986; Buresh et al., 1997). In plants, phosphorus serves as a structural element in nucleic acids and plays an important role in energy transfer and other enzyme processes (Marschner, 1995). Common diagnostic properties of phosphorus deficiency are a darker green leaf colour due to higher chlorophyll contents (often with red pigments from anthocyanins), reduced leaf extension and a higher root-to-shoot ratio, since root growth is much less affected by phosphorus deficiency than shoot growth (Wild, 1988; Marschner, 1995). A high phosphorus supply is needed for nodulation of legumes and hence phosphorus deficiency can also seriously reduce biological nitrogen fixation (Marschner, 1995). The minimum phosphorus concentration required for nodulation of soybean was about 0.5 mg P l–1 in external solution (Marschner, 1995). For some trees suitable for agroforestry, however, no enhanced biological nitrogen fixation was noted with phosphorus applications on phosphorus-deficient soils (Reddel et al., 1997; see also Section 13.1). In contrast to nitrogen, soil phosphorus is almost entirely derived from primary minerals, mainly apatite. Inorganic phosphorus is present in the soil solution as phosphate (H2PO4–/HPO42–). It adsorbs to clay mineral surfaces by ligand exchange and by specific bonding (bridging ligands which form two ligand exchanges) or precipitates with calcium and aluminium depending on the soil reaction (Mott, 1988). The pH for optimum phosphorus availability ranges from 5.5 to 6.5 when determined in water. The results of phosphorus analyses are greatly affected by the point of sampling, both vertically and horizontally. Inorganic phosphorus is very immobile in soil and small differences in sampling depth may yield completely different soil phosphorus contents. Vegetation effects (Tiessen et al., 1999) and application of organic or inorganic phosphorus sources to certain crops during previous cropping seasons (Woomer et al., 1998) cause uneven soil phosphorus distribution at a field and farm scale. This spatial variability creates challenges for scientific experimentation (Buresh, 1999).

Standard extraction procedures Extraction procedures for plant-available soil phosphorus use bicarbonate (Olsen), acid ammonium fluoride (Bray 1 and 2), hydrochloric and sulphuric acid (Mehlich 1, ‘double acid’), acetic acid and ammonium fluoride (Mehlich 2 and 3), ammonium carbonate and diethylenetriamine penta-acetic acid (DTPA) (Fixen and Grove, 1990), or anion exchange

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resins (van Raij et al., 1986). Inorganic phosphorus in the extract is most commonly analysed with the molybdate-ascorbic acid (molybdene-blue) procedure of Murphy and Riley (1962). Inorganic phosphorus yields may be affected by soil pretreatment: various studies have shown that air-drying of soil samples may lead to considerably higher (14–184%) extractable inorganic phosphorus due to liberation of phosphorus from microbial biomass (Sparling et al., 1985; Srivastava, 1998). Summaries of soil test methods for plant-available phosphorus and their interpretation have been provided by Olsen and Sommers (1982), Novais and Smyth (1999), Fixen and Grove (1990) and several chapters in Carter (1993).

Adsorption experiments In phosphorus-fixing soils, phosphorus fertilizer may be rapidly transformed into unavailable forms. The extent to which this happens depends on soil mineralogical properties (e.g. clay and especially allophane content and ferruginous nodules; Tiessen et al., 1991), but also on the history of phosphorus applications and plant uptake. Two adjacent fields may have the same available phosphorus contents measured with a standard procedure, but respond completely differently to mineral phosphorus applications because their adsorption sites for phosphorus are saturated to a different degree. Adsorption experiments are a means of predicting the magnitude of fixation from applied inorganic phosphorus. Soil is equilibrated with different solutions containing increasing phosphorus concentrations, and adsorption isotherms are determined from the relation between adsorbed and dissolved phosphorus (Fox and Kamprath, 1970). From the proportion of immobilized to added phosphorus, the fertilizer quantity that is needed to increase the concentration in solution to the desired level can be calculated (Bache and Williams, 1971; Holford, 1979; Haggar et al., 1991). By determining adsorption isotherms, it was shown that applications of organic matter decreased phosphorus adsorption due to competition for adsorption sites (Iyamuremye and Dick, 1996). Hue (1991) showed that organic acids decreased phosphorus sorption, but more so when phosphorus was added to the soil after the acids than when the phosphorus was added first. Accordingly, adsorption did not decrease if soluble phosphorus fertilizer (triple superphosphate) was added together with mulch on an Alfisol from East Africa (Nziguheba et al., 1998). The extent of the reduction of phosphorus sorption may depend not only on the type of organic acids (Hue, 1991), but also on the phosphorus content of the applied organic matter (Singh and Jones, 1976). Recent greenhouse studies have shown that high calcium contents of leaf mulch may decrease the solubility of phosphate rock. Consequently, phosphate rock did not increase maize


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growth when it was applied together with mulch (Smithson, 1999). In contrast, the dissolution of phosphate rock was increased by farmyard manure (Ikerra et al., 1994).

Sequential fractionation methods A central goal of some agroforestry practices is to increase nutrient efficiency by keeping applied nutrients in available form or to make recalcitrant nutrients available (Sanchez, 1995). Application of organic 0.5 g soil

Resin strip, 16 h

Resin Pi

0.5 M NaHCO3, 16 h

Bicarbonate Pi Bicarbonate Po

0.1 M NaOH, 16 h

Hydroxide Pi Hydroxide Po

1 M HCl, 16 h

Dilute Acid P

Conc. HCl, 10 min, 80˚C

Conc. HCl Pi Conc. HCl Po

Conc. H2SO4, H2O2

Residual P

Fig. 5.3. Flow chart of the sequential phosphorus extraction method (after Tiessen and Moir, 1993). Pi, inorganic phosphorus; Po, organic phosphorus; conc., concentrated.

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Table 5.1. Schematic comparison of plant availability and characteristics of soil phosphorus pools obtained by different analytical methods. Plant availability P pools

Soil solution extraction Bray, Olsen, Mehlich Isotopic exchange Microbial biomass Sequential extraction Physical fractionation NMRc

Not available

Slowly available

Readily available

Immediately available

Occluded inorganic and long-term mineralizable organic

Adsorbed inorganic and medium-term mineralizable organic

Exchangeable inorganic and short-term mineralizable organic

Soluble inorganic and rapidly mineralizable organic

(inorganic)a

inorganic inorganic organicb inorganic/organic inorganic/organic inorganic/organic

inorganic/organic inorganic/organic (inorganic/organic)

inorganic/organic inorganic/organic (inorganic/organic)

inorganic/organic inorganic inorganic organic inorganic/organic inorganic/organic

a

Brackets indicate less information about the respective pool from the applied method. Only part of short-term mineralizable organic P. c No differentiation according to plant availability possible; generally only NaOH-extractable P. NMR, nuclear magnetic resonance. b

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nutrient sources such as compost, manure or mulch and the transformation of recalcitrant nutrient pools into more readily available organic nutrient sources are key aspects of agroforestry. Organic nutrient management also implies that changes in available phosphorus will not be adequately assessed by conventional extraction methods for inorganic phosphorus. Mineralization and immobilization processes, and consequently the flow from inorganic to organic phosphorus pools and vice versa, become more important. Therefore, a single analysis of available phosphorus will very often not be sufficient to estimate the ability of a soil to supply phosphorus to plants. Sequential fractionation methods have been used for determining various soil organic and inorganic phosphorus pools with differing ecological properties. The Hedley fractionation method (Hedley et al., 1982) has been widely adopted and latterly further modified (Fig. 5.3). The basic concept of the method is to sequentially extract several soil phosphorus compounds from the same soil sample by using extractants with increasing strength. The fractions that are extracted first have high plant availability, whereas the fractions that are extracted last have very low plant availability (Table 5.1). Organic phosphorus is determined by subtraction of inorganic phosphorus (determined after precipitation of organic matter) from total phosphorus (determined in digested samples; Tiessen and Moir, 1993). Bicarbonate and hydroxide extractions can also be carried out on separate subsamples to avoid phosphorus losses during the sequential soil treatment (Nziguheba et al., 1998; Mutuo et al., 1999). Soluble organic phosphorus is usually not determined by resin extraction, but may be an important pool and can be obtained from soil solution extraction (see Section 7.2). Bowman et al. (1998) suggested a more rapid method for analysing occluded and resistant soil phosphorus by subtracting acid-extractable phosphorus in an ignited sample from total soil phosphorus. A single-step method for determining organic soil phosphorus using concentrated sulphuric acid was found suitable for Nigerian savanna soils (Agbenin et al., 1999). With sequential fractionation it has been possible to assess the distribution of applied phosphorus in the soil profile (Beck and Sanchez, 1996), the proportion of biological to geochemical phosphorus (Cross and Schlesinger, 1995), the soil incorporation of phosphorus from mulch or manure (Ikerra et al., 1994; Iyamuremye et al., 1996; Nziguheba et al., 1998; Solomon and Lehmann, 2000), the effects of burning (Ball-Coelho et al., 1993) and tree effects on soil organic phosphorus (Tiessen et al., 1992, 1999; Solomon and Lehmann, 2000; Lehmann et al., 2001c). Tree-specific phosphorus changes, especially in the dilute acid and organic phosphorus fractions, were found in a multistrata agroforestry system on an Oxisol in central Amazonia (Lehmann et al., 2001c), whereas rice monocropping, legume pasture and native savanna affected inorganic

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phosphorus in the bicarbonate and hydroxide pools of a Colombian Oxisol (Friesen et al., 1997). The inorganic as well as organic bicarbonate and hydroxide phosphorus fractions did not show fertilizer effects as clearly as resin phosphorus in a tropical Alfisol (Mutuo et al., 1999). An equilibrium between soil fractions seems to exist which replenishes plant-available resin-Pi from more recalcitrant phosphorus forms (bicarbonate and hydroxide) upon phosphorus uptake by plants (Schmidt et al., 1997). With phosphate rock applications, a combined anion–cation resin gives a better measure of phosphorus availability than an anion resin alone, because it binds soluble calcium, thereby increasing phosphate solubility (Saggar et al., 1992). In shifting cultivation in north-eastern Brazil, decline of phosphorus fertility resulted from mineralization of organic phosphorus and subsequent fixation to the mineral soil matrix rather than from net phosphorus export (Tiessen et al., 1992). A problem with phosphorus fractionation is that the ecological functions of the different fractions are not clear-cut and may depend on soil properties and management. The first two steps of the Hedley fractionation, resin and bicarbonate extraction, are standard methods for plant-available phosphorus (see above). However, the more recalcitrant fractions can be difficult to interpret. For example, the exchange between occluded and adsorbed phosphorus (characterized by acid-extractable and residual phosphorus) on the one hand and plant-available phosphorus on the other hand depends on their strength of adsorption. Also, the phosphorus supply capacity from organic sources may not be adequately assessed with sequential fractionation, as the activity of acid or alkaline phosphatase in soil may be more sensitive. In soils with high mineralization capacity, Oberson et al. (1996) found higher acid phosphatase activity after application of organic materials than in the control, whereas bicarbonate Po did not show any significant differences. A method for the direct measurement of phosphorus mineralization has recently been developed using phosphorus isotopes (Oehl et al., 2001), but has not yet been tested in tropical soils.

Microbial biomass phosphorus Next to nitrogen, phosphorus is the most abundant nutrient in microbial biomass. It constitutes a highly active pool with large turnover rates and short-term mineralizability (Brookes et al., 1984) (Table 5.1). The analysis follows the fumigation procedure using bicarbonate extraction as described by Brookes et al. (1982). The extraction of phosphorus from fumigated soil with strips of anion exchange membranes was found to be more rapid and accurate than that with bicarbonate, especially in strongly phosphorus-fixing soils (Saggar et al., 1990; Kuono et al., 1995). The soil


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water content prior to the extraction has an important impact on microbial phosphorus results and needs to be monitored (Sparling and West, 1989). Microbial phosphorus was found to be the second most sensitive indicator of soil phosphorus differences between intensive fallow and maize monoculture after phosphorus in the light fraction of soil organic matter (Maroko et al., 1999). Effects of applications of Tithonia diversifolia mulch on microbial soil phosphorus were inconsistent between different studies from western Kenya. Small effects of organic phosphorus applications may not be readily detected in soil microbial phosphorus (Jama et al., 2000).

Radioisotopes Adsorption experiments can also be conducted with 32P to determine isotopically exchangeable phosphorus (Fardeau et al., 1996; Frossard and Sinaj, 1997). Whereas green manure did not affect phosphorus extracted by Bray’s solutions or resin or phosphorus adsorption on an Oxisol in Brazil, it increased the amount of isotopically exchangeable phosphorus (LeMare et al., 1987). Exchange processes between organic and inorganic soil phosphorus pools have also been studied with the radioisotopes 32P and 33P (Di et al., 1997). The difference between exchangeable 32P on a non-incubated soil (only physicochemical exchange) and the 32P measured from incubated soils labelled with 32P (physicochemical exchange plus phosphorus mineralization) gives an estimate of phosphorus mineralization (Lopez-Hernandez et al., 1998). Isotopically labelled phosphorus fertilizer can also be used to follow its fate in soil phosphorus fractions (He and Zhu, 1997) or in the soil profile as affected by organic applications (Othieno, 1973). Additionally, the utilization of different soil phosphorus pools by different plant species may be estimated by 32P labelling of the soil: plants utilizing unlabelled phosphorus are able to access phosphorus pools that are more recalcitrant than the isotopically exchangeable, soluble phosphorus fraction as shown for lupin (Braum and Helmke, 1995). For valid interpretation of radioisotope experiments, care has to be taken to meet underlying assumptions of isotope exchange kinetics (Frossard and Sinaj, 1997).

Other methods From the total organic phosphorus in soil, no more than one-third has been chemically identified (Stevenson and Cole, 1999). Specific phosphorus compounds in soil comprise inositol phosphates, phospholipids, nucleic acids, phosphoproteins and metabolic phosphates

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such as ATP. Organic and inorganic bonding of soil phosphorus can be analysed by nuclear magnetic resonance (31P NMR) spectroscopy (Newman and Tate, 1980). With this method, increased amounts of labile organic phosphorus and, following mineralization, also of labile inorganic phosphorus were found after grassland conversion into conifer plantations (Condron et al., 1996). Relatively labile diester-P compounds were shown to accumulate in soil from earthworm casts (Guggenberger et al., 1996) and following application of manure (Solomon and Lehmann, 2000). A combination of physical fractionation of soil (see Section 4.3) with phosphorus analysis can increase options for assessing transformation processes which are not detected by analyses of phosphorus in the bulk soil. This approach may, in many cases, be more useful than analysing organic phosphorus fractions on bulk soils, because it provides information on turnover and mineralizability of soil organic matter and therefore also of associated organic phosphorus (Tiessen et al., 1999). Total phosphorus analyses, sequential phosphorus fractionation and 31P NMR spectroscopy have been used for phosphorus determination in density and particle size separates (Maroko et al., 1999; Solomon and Lehmann, 2000). Maroko et al. (1999) found that phosphorus effects of planted Sesbania sesban fallows and natural fallows, in comparison with continuous maize cropping, were most evident in the light organic matter fraction compared with a range of other phosphorus analyses. Soil phosphorus availability for plants is influenced by the presence and activity of phosphorus-solubilizing bacteria in the rhizosphere, root release of protons and organic acids, mycorrhizal root infection, and root hair length and density. These factors are more important for the uptake of phosphorus than for that of any other macronutrient due to the low mobility of phosphorus in soil and its strong fixation to oxides. The ultimate test for soil phosphorus availability is plant uptake. Bioassays can give valuable information about the effects of different management interventions such as applications of mulch or mineral fertilizer on plant phosphorus uptake (Othieno, 1973; Haggar et al., 1991; Jama et al., 2000). Phosphorus transfers in soil, e.g. from topsoil to subsoil, are seldom determined due to experimental problems of obtaining phosphorus from soil solution with ceramic suction cups or lysimeters (see Section 7.2). With little fertilization, amounts of inorganic phosphorus are usually low and possess low mobility especially in high phosphorus-fixing soil. In the absence of soil erosion, phosphate losses from agricultural soils do not usually exceed 0.5 kg ha–1 year–1 (Sharpley and Withers, 1994). However, organic phosphorus can be the dominant phosphorus compound in soil solution and dissolved organic carbon properties largely determine phosphorus mobility in soil (Qualls et al., 1991; Donald et al., 1993). Donald et al. (1993) demonstrated for a forest soil that 64% of the organic phosphorus was contained in the hydrophobic neutral fraction of dissolved


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organic carbon that was only weakly adsorbed to the soil. Therefore, soil solution phosphorus can be mobile in its organic form (Schoenau and Bettany, 1987) and should be considered when calculating phosphorus budgets.

5.4 Methods for Soil Sulphur ( J. Lehmann) Arable soils of the humid tropics often have low total sulphur contents because of low contents of parent materials, strong weathering and high leaching losses. In comparison with other macronutrients, sulphur received little attention as a plant nutrient in tropical crop production in the past and few results have been published (Kang et al., 1981; Acquaye and Kang, 1987; Motavalli et al., 1993). Sulphur deficiency in tropical agroecosystems has recently increased, however, due to the more common use of nitrogen and phosphorus fertilizers with low sulphur contents (e.g. substitution of ammonium sulphate by urea and of single by triple superphosphate) as well as higher crop yields in some regions (Ceccotti et al., 1998). Up to 90% of sulphur in plants is present as amino acids and therefore bound in proteins. Sulphur is an essential element for many functions in plants (e.g. synthesis of chlorophyll) and sulphur deficiency not only decreases plant growth but also forage quality (Tisdale, 1977) and resistance to pests and diseases. Visual diagnosis of sulphur deficiency in plants is difficult and criteria range from chlorosis in young leaves, starting from the edges, spoonlike deformations, succulence, poor pod formation, to stunted growth (Schnug and Haneklaus, 1998). Deficiency symptoms are more difficult to diagnose for monocotyledons than for dicotyledons and resemble those of nitrogen. Tissue analysis of total sulphur is a more reliable indicator of sulphur deficiency than visual diagnosis if the usual sampling criteria are followed, such as the collection of defined plant organs and development stages (Schnug and Haneklaus, 1998).

Sulphate-S Plants take up sulphur in the form of SO42–, which is the most common form of inorganic sulphur in agricultural soils. Grasses have a higher ability than legumes to utilize sulphate in soils (Havlin et al., 1999), and excessive competition for sulphate-S in legume–non-legume mixtures may cause decreases in biological nitrogen fixation. Common extractants for plantavailable sulphate-S include water, calcium chloride, sodium phosphate (Na2HPO4) (Kowalenko, 1993a), Na-acetate/acetic acid (Landon, 1991), lithium chloride (Tabatabai, 1982) and exchange resins (Searle, 1992).

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However, the value of analysing sulphate-S for estimating the availability of soil sulphur for plants is not entirely clear (Schnug and Haneklaus, 1998) and additional availability tests are recommended. Sulphate sorption characteristics are important for quantifying sulphur leaching and retention in tropical soils. Their determination is analogous to phosphorus sorption experiments (see Section 5.3). In soils high in clay, organic matter and exchangeable acidity, soil drying prior to analysis can result in large errors and experimentation with field moist samples is recommended (Comfort et al., 1991). The extent of sulphate sorption in acid subsoils can differ from that of the topsoils as shown for an Alfisol and Oxisol from India (Patil et al., 1989), but recycling by deeprooting trees as occurs with nitrate (Box 8.1) has not yet been the subject of research. Adsorption generally increases with higher clay contents, oxide contents, and lower pH. Adsorption and precipitation of sulphateS by carbonates has to be considered in certain dryland soils.

Mineralization of sulphur from soil organic matter and biomass More than 95% of sulphur in soil occurs in an organic form. Since organic sulphur cannot be determined directly, total sulphur is usually measured using dry combustion and gas chromatography or dry ashing with alkaline oxidation followed by ion chromatography (Tabatabai et al., 1988). Organic sulphur is calculated as the difference between total and inorganic sulphur. Sulphur availability for plants is closely linked to the mineralization of organic sulphur, similar to soil nitrogen (see Section 5.2). Sulphur mineralization can be measured by incubation methods. However, much higher mineralization rates of sulphur (but not nitrogen) have been measured when the mineralized sulphur was periodically leached out of the soil than in closed systems, indicating that certain incubation methods that are commonly used for nitrogen would lead to underestimation of sulphur mineralization rates (Maynard et al., 1983). Therefore, mineralization studies are best conducted in open systems under field conditions (Eriksen et al., 1998), where leaching of sulphate-S is allowed and mineralization processes are controlled by ambient temperature and moisture conditions. Plants have been shown to stimulate sulphur mineralization in the soil through a greater proliferation of microorganisms in the rhizosphere (Stevenson and Cole, 1999), which can make the prediction of sulphur availability more difficult when estimated from mineralization studies where roots are excluded. The effect of sulphur from organic sources such as leaf mulch, animal manure or compost on sulphur availability depends on the concentration of sulphur in the organic materials. Sulphur immobilization in microbial biomass occurred after application of barley straw with low sulphur content


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(2.2 mg g–1), but not of rape leaves with high sulphur content (7.2 mg g–1; Wu et al., 1993). However, only a rough threshold value can be given for sulphur release from organic residues, which tend to mobilize sulphur if the C:S ratio is 400 (Stevenson and Cole, 1999). Sulphur release was found to be quite different from that of nitrogen, as soluble sulphate-S is initially solubilized in high amounts and sulphur mineralization is not closely related to the C:S ratio of the organic material (Janzen and Ellert, 1998). Short-term increases of available sulphur from applied animal manure can rarely be expected, although long-term accumulation of manure or urine in pasture soils increases soil organic sulphur (Saggar et al., 1998) and leads to a higher supply of sulphate-S.

Fractionation of soil organic sulphur Another approach to determining soil organic sulphur pools with different mineralizability is chemical fractionation. Ester-sulphate-S compounds can be analysed using reduction with hydriodic acid (HI) and subtracting inorganic sulphate-S. Carbon-bonded sulphur is then obtained by subtraction of ester-S from organic sulphur (Fig. 5.4). HI-reducible estersulphate-S is considered to be more labile than carbon-bonded sulphur, although some authors have reported the opposite (Janzen and Ellert, 1998). Carbon-bonded sulphur was not affected by trees in a multistrata agroforestry system in the central Amazon, whereas ester-sulphate-S increased in soil underneath trees with large nutrient recycling (Lehmann et al., 2001c). Solomon et al. (2001) reported that forest clearing and maize cultivation in the Ethiopian highlands reduced both the carbon-bonded and ester-sulphate-S, whereas, in plantations of tea and Cupressus lusitanica, only the carbon-bonded sulphur fraction decreased. In a simplified conceptual model, carbon-bonded sulphur is considered to be released by mineralization, which is controlled by the needs of soil microorganisms for organic carbon, whereas ester-sulphate-S is released by enzymatic hydrolysis and therefore controlled by the supply of sulphur (McGill and Cole, 1981; see also comments in Ghani et al., 1992). This dichotomous cycling of sulphur also explains variable N:S ratios of mineralized organic matter. Usually, sulphur was found to be more stable in organic matter than nitrogen upon soil cultivation (Stevenson and Cole, 1999). The dynamics of sulphur fractions under long-term cultivation have also been studied in humin, humic and fulvic acids (Bettany et al., 1980); however, the analysis of ester-sulphate-S and carbon-bonded sulphur in these extracts may be affected by hydrolysis of organic sulphur compounds during the extraction and may not yield reliable results (Freney, 1986). The analysis of organic sulphur in soil physical fractions offers the

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0.5 g soil

0.5 g soil

0.01 M CaCl2, 30 min drying at 100˚C

0.5 g soil

Na-hypobromite, 3 min drying at 250˚C

Distillation with HI acid



Inorganic sulphate-S

Total HI-reducible S

Total S

Calculations: Total S – Inorganic sulphate-S = Organic S Total HI-reducible S – Inorganic sulphate-S = Ester-sulphate-S Organic S – Ester-sulphate-S = Carbon-bonded S

Fig. 5.4. Conceptual outline of the sulphur fractionation according to Kowalenko (1993b). Total sulphur can also be obtained by dry combustion with an automated analyser.

possibility to detect tree effects on soil sulphur more sensitively and also to evaluate its stability after applications of mulch or animal manure. Sulphur can be analysed after fractionation according to particle or aggregate size and density (Anderson et al., 1981). A fractionation method has been described which uses acetylacetone as extractant and different intensities of ultrasonic dispersion (Eriksen et al., 1995). Microbial sulphur comprises a pool of highly reactive sulphur in soils and is important for understanding the mineralization of carbon-bonded sulphur, although it only amounts to 1–2.5% of total soil sulphur. Microbial sulphur can be determined by the fumigation–extraction method using calcium chloride extraction (Saggar et al., 1981; Wu et al., 1994). Other soil organic sulphur measurements such as analyses of enzymes, amino acids and sulpholipids have been developed, but have not yet yielded easily interpretable results.

Other methods Isotopic techniques using radioactive 35S have been successfully employed to identify the fate of added sulphur in different soil pools and its subsequent mineralization (Freney et al., 1971; Maynard et al., 1983; Ghani et al., 1993). Soil is incubated with carrier-free 35SO42– in the presence or


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absence of carbon sources (glucose) and sulphate, and analysed for 35S. Excess sulphate is leached and the soil is re-incubated. With this method, processes such as the incorporation of added sulphur from mulch or animal manure and the incorporation into soil sulphur pools such as estersulphate-S, carbon-bonded sulphur and sulphate-S can be studied. When studying the sulphur balance of agricultural systems, exchanges of sulphur between the vegetation and the atmosphere have to be considered (Krouse et al., 1991). If atmospheric sulphur concentrations are high, such as near the sea, sulphur uptake by plant leaves from the air can sometimes occur in sufficient amounts to meet plant requirements. On the other hand, plants may also release volatile sulphur compounds in response to nutritional or light stress. Sulphur losses from decomposing litter can be greater than from living plants (Janzen and Ellert, 1998).

5.5 Methods for Potassium, Calcium and Magnesium in Soil (G. Schroth, J. Lehmann) The cations potassium, calcium and magnesium occur in several forms in the soil that differ in their availability to plants. The most readily plantavailable fraction is that in the soil solution, followed by the exchangeable fraction, which replenishes the soil solution if nutrients are removed by either plant uptake or leaching. Potassium fixed in clay interlayers becomes available at a time scale from hours to weeks. The least available forms are various primary and secondary soil minerals, which release the respective nutrients upon weathering (Haby et al., 1990). Together with sodium (where this is present at relevant quantities), potassium, calcium and magnesium make up the exchangeable bases of a soil. Numerous soil test methods are used for assessing the availability of these nutrients to plants, which are discussed in detail by Haby et al. (1990). As a rule, those methods that have been found most useful in the respective study region and for which locally calibrated reference values exist should be used. Of particular interest are multielement extractants such as Mehlich 3, which allows the simultaneous extraction of the exchangeable bases, phosphorus and various micronutrients (Tran and Simard, 1993). Acid tropical soils, such as Oxisols and Ultisols, are characterized by low base saturation, and correspondingly low exchangeable contents of potassium, calcium and magnesium. Despite low contents, potassium availability rarely limits crop yields when acid soils are first taken into cultivation, although it becomes an important factor for permanent, intensive cropping once these soils have been limed and other deficiencies (e.g. phosphorus) have been corrected (von Uexküll, 1986). Small quantities of lime often increase potassium uptake by plants due to improved root growth, whereas larger quantities may depress potassium

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uptake by increasing the cation exchange capacity associated with variable (pH-dependent) charges in the soil, thereby reducing potassium concentrations in the soil solution, and by antagonistic effects of calcium ions on potassium uptake. Low magnesium contents are common in acid soils, and magnesium deficiency can be induced by high aluminium contents (see Section 5.6) or by potassium fertilization. Tree crops tend to be more sensitive to magnesium deficiency than annual crops, and annual dicotyledons more sensitive than annual monocotyledons (von Uexküll, 1986). With time under cropping the exchangeable soil contents especially of calcium and magnesium decrease and exchangeable acidity increases, unless cation losses caused by the exportation of harvested products, leaching and erosion are replaced by corresponding inputs of fertilizers and lime (Pieri, 1989; Juo et al., 1995; Smyth and Cassel, 1995). In most soils, calcium and magnesium are more susceptible to leaching than potassium (see Section 7.1). If land is left to regenerate under spontaneous or planted fallow vegetation, losses of calcium (and probably other cations) in the plant–soil system may continue during the first years before nutrients start to accumulate due to atmospheric inputs, mineral weathering (where weatherable minerals are still present) and uptake from the subsoil (Szott et al., 1999). Tree species differ in their effects on nutrient cations in the soil, and such species-related differences can even be detected within closed forest stands (Finzi et al., 1998). Trees are expected to reduce nutrient leaching from cropping systems (see Chapter 7), but they may also immobilize relevant quantities of nutrients in their biomass. In a long-term experiment, Hulugalle (1994) found lower exchangeable calcium contents in the soil under hedgerow intercropping than annual crops and pasture and explained this in terms of high calcium demand of the trees. Larger amounts of calcium and magnesium were immobilized in unpruned Cordia alliodora than in pruned Erythrina poeppigiana shade trees in coffee plantations (Beer et al., 1998), and immobilization of potassium in the Cordia alliodora biomass was suspected to be a potential limiting factor to crop and tree productivity (Beer, 1988). Some trees promote the accumulation of cations in the topsoil, presumably through efficient uptake by a large root system and subsequent release from litter. Reported cases include calcium accumulation by Gmelina arborea (Sanchez et al., 1985; Fisher, 1995) and Senna siamea (Drechsel et al., 1991). Elevated calcium contents also characterize the leaf litter of Terminalia superba (F. Bernhard-Reversat, unpublished data). Several tree species increased the calcium and potassium contents in the topsoil of a degraded rainforest site (Fisher, 1995). Relatively high potassium concentrations are found in the biomass of the large herbaceous plant, tithonia (Tithonia diversifolia), which is also rich in nitrogen and


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phosphorus. The effectiveness with which this potassium was used by maize when supplied with tithonia biomass seemed to be comparable to the use of potassium from mineral fertilizer (Jama et al., 2000). Increased potassium contents in the topsoil under certain cover crops have been explained with potassium redistribution from lower soil horizons (Smyth et al., 1991; Barber and Navarro, 1994). In accord with their different functions in the plant, potassium and calcium differ strongly in their release dynamics from both dead and living biomass. Potassium is rapidly leached from leaf and woody litter, whereas calcium often shows an absolute increase in biomass during the first stages of decomposition, possibly due to a combination of slow release and calcium import in fungal hyphae (Swift et al., 1981; Maheswaran and Gunatilleke, 1988; Schroth et al., 1992). Potassium is also leached in relatively large quantities from living plants. In studies in a rainforest in Borneo (Burghouts et al., 1998) and in multistrata agroforestry and fallow in Amazonia (Schroth et al., 2001a), potassium was the only macronutrient for which larger quantities were recycled from the standing biomass to the soil in throughfall and stemflow than in litterfall. Cycling of calcium, magnesium and potassium in agroforestry systems can also be studied with tracers. Due to handling difficulties in conjunction with health hazards, radioisotopes (42K, 28Mg, 45Ca) are only used in specialized experimentation under controlled conditions. However, subsurface applications of the elements lithium and rubidium (for potassium) and strontium (for calcium) can be used to assess root activity distribution (Fitter, 1986; van Rees and Comerford, 1986) (see Section 8.2). The determination of the fate of applied potassium and calcium fertilizer with these tracers may pose difficulties, because they would have to be applied at high concentrations, which would alter their behaviour in soil and during plant uptake.

5.6 Methods for Soil Acidity (G. Schroth) Acid soils, especially Oxisols and Ultisols, occupy vast areas in the tropics. Naturally, these soils are mostly covered by forest or savanna, but during the last decades increasing areas have been cleared for agriculture. Despite the acidity and nutrient deficiency of these soils, their economic importance is substantial: 100% of the world tea, rubber, oil palm and cassava production and 90% of the world coffee production come from acid soils (von Uexküll and Mutert, 1995). However, the conversion of forest on acid soils for agricultural use very often leads to the development of unproductive, man-made savannas, such as the imperata grasslands of South-east Asia and some abandoned pastures of Amazonia. In the absence of the necessary inputs of nutrients and lime, the residual fertility from

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the clearing of the vegetation is lost after only a few crop harvests, and the land is then commonly abandoned. Under conditions of increasing pressure on acid soils in the tropics from a growing population, the problems of soil acidity and infertility are intimately connected to losses in tropical forest area and biodiversity (von Uexküll and Mutert, 1995). Acid soils require a humid climate for their development and are thus typically found in tropical savanna and rainforest zones. Under agricultural use, soil acidification is increased by leaching of basic cations and their removal with harvested crops; acidifying fertilizers such as urea, ammonium sulphate and potassium chloride; and the use of nitrogenfixing legumes, for example, in pastures (Rowell, 1988; von Uexküll and Mutert, 1995). Agroforestry techniques may reduce problems of soil acidity by reducing leaching losses (see Chapter 7) and by increasing the level of soil organic matter, which detoxifies dissolved aluminium (see Chapter 4), although increased acidification due to the use of nitrogen-fixing trees may also be expected (van Miegroet and Cole, 1984). Acid soils present a number of interrelated problems, including toxicity of aluminium, manganese and (under reducing conditions) iron as well as deficiencies of phosphorus, calcium, magnesium, potassium and micronutrients (e.g. molybdenum, boron). Low water-holding capacity (e.g. of Oxisols) and susceptibility to crusting, erosion and especially compaction (Oxisols, Ultisols) aggravate the problem (Marschner, 1995; von Uexküll and Mutert, 1995). Here, only aluminium toxicity as a key factor of acidity stress in many mineral soils is discussed. The risk of manganese toxicity is lower than that of aluminium toxicity in highly weathered tropical soils, because these often have low total manganese contents (Marschner, 1995). Mechanisms of plant adaptation to soil acidity have been discussed by Marschner (1995) and, specifically for tree species, by Fisher and Juo (1995). Lists of acid-tolerant and intolerant annual and perennial crop and pasture species and varieties are provided by Sanchez and Salinas (1981), and much useful information on tree species for acid soils can be found in the volume edited by Evans and Szott (1995).

Diagnosis of aluminium toxicity in plants In soils with a pH in H2O >5.5, aluminium is tightly held to the exchange surfaces, but as the pH falls below 5, exchangeable aluminium and aluminium concentrations in the soil solution increase markedly and can cause aluminium toxicity in sensitive plant species (von Uexküll, 1986). Neither visual diagnosis of shoots nor foliar analysis for aluminium is a suitable indicator of aluminium stress in cultivated plant species. Most of these are aluminium excluders and do not readily transport aluminium from the roots into the shoots, a notable exception being tea (Camellia


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sinensis). In fact, aluminium concentrations in shoots of healthy plants may be higher than in those of aluminium-stressed plants. In tropical rainforests, aluminium includers and excluders exist at the same site, and these may vary in their foliar aluminium contents by two orders of magnitude. Aluminium concentrations in roots of aluminium-stressed plants are correlated with the severity of damage, but the analysis of roots for aluminium is difficult due to adhering mineral soil (Bergmann, 1988; Marschner, 1995). Symptoms of aluminium toxicity first appear on the roots, which are shortened, thickened, show reduced branching and are brown to black in colour with black tips. Symptoms are limited to the actively growing tissue (Bergmann, 1988). Reduced root development, especially in the subsoil, makes plants sensitive to drought stress and reduces the access to nutrients in the subsoil, thereby also increasing nutrient leaching (Rowell, 1988). High aluminium concentrations in the solution impede the uptake of polyvalent cations such as calcium and magnesium and may thus induce deficiencies of these elements, whereas potassium uptake is usually unaffected (Marschner, 1995). Aluminium interacts with phosphorus availability through the reduction of root growth and by binding phosphate on root surfaces, cell walls and in the free space of plant roots. At higher aluminium concentrations, symptoms of aluminium toxicity may, therefore, be similar to those of phosphorus or calcium deficiency (Bergmann, 1988; Rowell, 1988).

Aluminium measurement in soil To relate problems of aluminium toxicity and induced nutrient deficiencies in plants to soil properties, aluminium may either be measured in the dry soil or in the soil solution. The former approach is more compatible with standard soil sampling and laboratory practices and is, therefore, more common. Exchangeable acidity is usually measured by leaching a soil sample with 1 M KCl and quantification of the extracted acidity (aluminium and protons) by titration (Hendershot et al., 1993b). Exchangeable acidity is given in mmolc kg–1 and/or as a percentage of the effective cation exchange capacity measured at the pH of the soil, e.g. by summing the exchangeable cations (Hendershot et al., 1993a). In relatively aluminium-tolerant crop species and varieties, toxicity symptoms may appear at 60% aluminium saturation, and in less tolerant species and varieties at 30% aluminium saturation (von Uexküll, 1986). A procedure for fractionating aluminium forms in soils has been proposed by Soon (1993). However, plants are directly influenced by aluminium in the soil solution, which is not always proportional to exchangeable aluminium. In

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a comparison of the soil and soil solution chemistry of different land-use systems in central Amazonia, fallow and low-input agroforestry had more exchangeable acidity than high-input agroforestry and monoculture plantations, but the high-input systems had more aluminium in the soil solution due to the increased availability of fertilizer cations for exchange reactions with the sorbed aluminium and the acidifying effect of nitrification of fertilizer nitrogen. Peak concentrations of aluminium in the soil solution coincided with periods of highest nutrient availability following fertilizer applications and could have negatively affected the uptake of fertilizer nutrients (Schroth et al., 2000b). Critical aluminium concentrations in the soil solution which negatively affect plant growth vary from about 1 mM for tolerant species to 1 mM for sensitive species (Rowell, 1988). In legumes, the critical concentration for nodulation may be substantially lower than that for the host plant (Ashwath et al., 1995; Marschner, 1995). The principal difficulty in interpreting aluminium concentrations in soil solution (but also exchangeable aluminium levels) is that, depending on the solution composition, different aluminium forms coexist which may differ widely in their phytotoxicity (Kinraide, 1991). Organic substances such as fulvic acids, humic acids and other organic acids may detoxify aluminium by forming complexes. Consequently, aluminium in the subsoil can be phytotoxic at lower concentrations in the soil solution than in the topsoil where organic matter contents are higher (Marschner, 1995), and the application of mulch and green manure can reduce aluminium toxicity in acid soils (Harper et al., 1995). The concentration of different aluminium species in the soil solution can be predicted with models such as GEOCHEM, although not all the required stability constants may be accurately known (Kerven et al., 1995). Also, the actual solution composition can differ from that predicted by equilibrium models during peak flow conditions after heavy rain when aluminium release may be influenced by diffusion processes (Franken et al., 1995). Because of the suppressive effect of high aluminium concentrations on calcium and magnesium uptake, the molar Ca:Al and Mg:Al ratios in soil solution have been used as measures of aluminium-induced deficiencies of these elements and were superior in this respect to the concentrations of the individual elements (Marschner, 1995). In addition to the effects of soil acidity on plants via toxicity and induced deficiency phenomena discussed above, soil acidity may also influence plant growth by affecting soil biological properties such as the abundance and composition of the soil fauna (Lavelle et al., 1995) and plant diseases. Methods of determining lime requirements for crops of differing sensitivity to soil acidity have been discussed by Sumner (1997) and van Raij and Quaggio (1997). For the assessment of plant litter effects on soil acidity, see Chapter 6.

Chapter 6 Decomposition and Nutrient Supply from Biomass G. SCHROTH Biological Dynamics of Forest Fragments Project, National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil

6.1 Synopsis One of the benefits that may be expected from agroforestry in comparison with annual and perennial monocultures is an increased production of biomass. When returned to the soil, this biomass protects the soil surface, releases nutrients, replenishes soil organic matter and provides carbon substrates for soil biota. This chapter focuses on above-ground biomass, which is easier to manage and which has been studied more intensively than root biomass in agroforestry, but the principles are also valid for the latter. Methods for studying root turnover as an important source of biomass input in the soil are discussed in Section 12.4. With respect to nutrient inputs into the soil, there is a major difference between biomass grown in situ and fertilizers in so far as only part of the nutrients released from the biomass will be an external input, the remainder has been taken up from the same soil to which it is returned. Biomass transfer systems represent an intermediate situation in which the nutrients in the biomass are added to the site where the biomass is applied, but are removed from another site within the same landscape. Net nutrient additions in biomass to a system include biologically fixed nitrogen, nutrients taken up by the trees either from deeper subsoil horizons or from recalcitrant pools that would not have been readily available to crops. Nutrients taken up by extensive lateral tree roots beyond field boundaries are again an intermediate case. They can be seen as net additions to the system if they are taken up from a site that would have never been cropped (a river bed), but only as redistribution if they are taken up from a © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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neighbouring field or from a fallow that will be cropped the following year, depending on the spatial and temporal scale under consideration. Other nutrients in biomass may not be additions, but rather ‘avoided losses’, such as nutrients taken up from the percolating soil solution that would have been leached in the absence of the trees (see Chapter 7). However, a significant proportion of the nutrients in tree biomass may have been taken up in direct competition with crops. If tree biomass is applied to the soil, these nutrients are merely recycled. In simultaneous agroforestry systems, this may often be the main part of the nutrients contained in tree biomass. In contrast, in fallows, where direct tree–crop competition for nutrients does not occur or only between neighbouring crop and fallow plots, much of the nutrients in the tree biomass may consist of net additions and avoided losses. The basic function of fallows is to produce biomass and thereby regenerate soil fertility. In the most widespread simultaneous agroforestry systems, the production of nutrient-rich biomass may also be an important function of the trees (e.g. pruned legume trees as shade in coffee and cocoa plantations), but it is usually not their only function and may often be of secondary importance to farmers in comparison with direct production functions (e.g. fruit or fodder trees in parkland systems in savannas). In hedgerow intercropping, the trees serve principally for the production of biomass and the recycling of nutrients through this biomass, but do not have direct production functions unless they are simultaneously used for fodder. The fact that this technique has not been widely adopted may indicate that these motives alone often do not provide sufficient incentive for farmers to plant and manage trees (Fujisaka, 1993). Agroforestry interventions need to be designed in the context of the niche within a farming system that the trees will fill. The optimization of carbon and nutrient supply will generally be constrained by farmer requirements for economic product, low competitiveness with crops and ease of management.

Nutrient cycling in biomass in agroforestry systems The quantities of tree litter or prunings that are produced in agroforestry systems and the amounts of nutrients therein can be substantial. In coffee and cocoa plantations shaded by legume trees in Central America, 3–14 Mg ha–1 year–1 of prunings are produced by the trees, containing 60–340 kg of nitrogen (Beer et al., 1998). Leguminous trees in hedgerow intercropping produced up to 20 Mg ha–1 year–1 of prunings, containing as much as 358 kg of nitrogen, 28 kg of phosphorus, 232 kg of potassium, 144 kg of calcium and 60 kg of magnesium (Palm, 1995). Very little is known about the amounts of carbon and nutrients released by root systems in agroforestry (see Section 12.4). Relevant nutrient release from roots

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may occur when large amounts of superficial tree roots are destroyed by tillage at the beginning of a cropping season, and following the conversion of forest or fallow plots into crop fields. The amount of biomass produced by agroforestry trees depends on the tree species, the number of trees per hectare, tree age, tree management and site factors. The nutrient concentrations in different parts of the biomass depend mainly on tree species, phenological stage (e.g. senescence, fruiting), management and site factors. Nitrogen-fixing trees normally have higher nitrogen concentrations in the biomass than non-fixing species, but this characteristic also varies widely between species (Palm, 1995). Deciduous species generally have higher nitrogen concentrations in the leaves than evergreen species (Eamus, 1999). Certain monocots such as bananas, bamboos and palms have relatively high potassium concentrations in their biomass (Sanchez et al., 1985). Tree prunings normally have higher concentrations of mobile nutrients such as nitrogen, phosphorus, potassium and zinc than naturally fallen litter from which these elements are retranslocated by the tree prior to abcission (Marschner, 1995). Similarly, the young, leaf-rich biomass of frequently pruned trees has higher nutrient concentrations than the more woody biomass of infrequently pruned trees, although the quantity of biomass produced decreases with pruning frequency (Duguma et al., 1988). Roots cut off during soil tillage would also be expected to have higher contents of certain nutrients than roots that die naturally, although the question of nutrient retranslocation from senescing roots requires further clarification (see Section 12.4). Trees will also have higher nutrient concentrations in their biomass when they grow in nutrient-rich than in nutrient-poor soil (Budelman, 1989; Palm, 1995). Compilations of nutrient concentrations in tree biomass can be found in Young (1997), Palm (1995), books on animal nutrition and in the Organic Resource Database developed by the Tropical Soil Biology and Fertility Programme (TSBF) and Wye College, London (www.wye.ac.uk/BioSciences/soil). The amount of nutrients contained in tree prunings has often been compared with the nutrient requirements of the crops to which the prunings were applied. Such comparisons are useful to illustrate the relative importance of the nutrient fluxes through the tree biomass, but it must be kept in mind that in simultaneous agroforestry systems a large part of the added nutrients might have also been available to the crops in the absence of the trees. According to Palm (1995), the nutrients in 4 Mg ha–1 of leaves from any of four leguminous tree species could meet the requirements of a maize crop for nitrogen, calcium and much of the magnesium and potassium. Nitrogen was not supplied in sufficient quantity to meet crop demands by two non-leguminous trees, and phosphorus was not supplied in sufficient quantity by any of the tree species studied. This latter point is also valid for other annual crops

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(Schroth et al., 1995c). Biomass is usually a less efficient source of phosphorus than of other macronutrients.

Synchrony and synlocation of nutrient release with plant uptake When biomass decomposes on or in the soil, the nutrients may either remain in the soil in mineral form, be incorporated in the soil biomass and soil organic matter (immobilization), be taken up by plants, or be lost from the system through leaching or in gaseous form. The relative importance of these different pathways depends on the respective nutrient, the decomposing material, and the biotic and abiotic conditions under which the decomposition process takes place. It has been hypothesized that the efficiency of the uptake of nutrients released from biomass by crops can be increased by improving the synchrony and synlocation (i.e. the temporal and spatial correspondence) of nutrient release with plant uptake. This concept can be applied to nutrient release both from biomass and from soil organic matter and is particularly relevant where losses of released nutrients are likely to be high, such as in humid tropical regions with high risk of leaching and denitrification during most of the year, or in savanna regions with a pronounced flush of nitrogen mineralization at the beginning of the rainy season (Myers et al., 1994; Heal et al., 1997). Nutrient release from biomass and soil organic matter, and thus its synchrony and synlocation with crop uptake, can be influenced by management. Nutrient release from biomass can be controlled by selecting plant materials with desirable nutrient release kinetics and adjusting the timing and method of their application, with nutrient release being faster from soil-incorporated than from surface-applied materials. Nutrient release from soil organic matter is affected by the method and timing of soil tillage, mulching to reduce wetting and drying cycles, and the use of cover crops. To be effective in improving the uptake of a limiting nutrient by crops, it is important to consider whether synchrony and synlocation of release either directly from tree biomass or from soil organic matter is most relevant, so that appropriate management measures can be designed. This underlines the importance of studies of the fate of nutrients following their release from biomass. When litter or mulch is exposed to rainfall on the soil surface, a soluble nutrient fraction is rapidly leached from the biomass into the soil either in mineral or organic form. This fraction includes most of the potassium, and for freshly cut (as opposed to naturally fallen) materials it may also include a relevant percentage of the phosphorus in the biomass (Babbar and Ewel, 1989; Schroth et al., 1992). Rapid initial release of nitrogen from Cajanus cajan prunings has been attributed to leaching (Schroth et al., 1992), but this has not been confirmed for other materials (Vanlauwe et


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al., 1995). The fate of such rapidly released nutrients, and especially whether they are taken up by crops or washed further downward in the soil, should depend on the presence and activity of root systems in the soil at this time and on pedoclimatic conditions. Up to now such studies tracing the fate of released nutrients have concentrated largely on nitrogen, and little information is available on what happens to other nutrients including, for example, the relatively large quantities of potassium leached from tree litter or prunings. The release of nitrogen from biomass and its fate in the soil–crop system has received considerable attention, both because of its importance to plant growth in many agroecosystems and the availability of suitable tracers which facilitate such research. Data compiled by Palm (1995) show that the recovery of biomass nitrogen by the crops to which it is applied is generally less than 20% and often closer to 10%. This compares with values of 50–70% for mineral fertilizers, although fertilizer use efficiency in Africa is often much lower, and 20–30% for manure (Finck, 1992; Giller and Cadisch, 1995). Studies with 15N-labelled legume biomass have shown that most of the nitrogen not recovered in the crops is incorporated into soil organic matter, from which it is then successively released through mineralization (Ladd, 1981). Accordingly, additions of legume tree prunings during 8 years of hedgerow intercropping caused substantial increases in soil nitrogen mineralization rates in Costa Rica (Haggar et al., 1993), and nitrogen mineralization rates in the soil were also substantially increased under a leguminous cover crop in an Amazonian agroforestry system (Schroth et al., 2001c). Such results suggest that the nitrogen benefit to crops from biomass comes mainly from the longer-term build-up of readily mineralizable soil organic matter pools rather than from the shortterm release of nitrogen from the decomposing material (Myers et al., 1994; Palm, 1995). Consequently, it may be more important to improve synchrony and synlocation of nitrogen uptake by plants with its mineralization from soil organic matter than with its release from biomass, and this should be reflected by research priorities (see also Section 5.2). Little is known about the relative importance of short-term release as opposed to incorporation into mineralizable soil organic matter pools for phosphorus and sulphur (see Sections 5.3 and 5.4).

Prediction of nutrient release from biomass As noted before, the dynamics of nutrient release from decomposing biomass under agroforestry conditions and the factors influencing it are the object of intensive studies. The principal objective of this research is to identify parameters that allow quantitative prediction of the time course of nutrient release from biomass as influenced by attributes of the biomass

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itself (the resource quality) and the physicochemical environment in which it is decomposing. As a rule, the decomposition of organic materials in soil leads to an initial net mineralization of nitrogen if the C:N ratio is 30. During the decomposition process, carbon is released as carbon dioxide, and the C:N ratio is progressively reduced below the threshold value for net mineralization. For phosphorus and sulphur, the corresponding threshold values are C:P400 for initial net immobilization. These values apply if carbon and the respective nutrient are in compounds with similar degradation rates (Stevenson and Cole, 1999). For the prediction of nitrogen release from litter and prunings of agroforestry tree species, the C:N ratio alone has not been found satisfactory, as several other chemical characteristics of biomass affect the decomposition dynamics, especially their lignin and polyphenol content (Mafongoya et al., 1998). Lignin, which is a low-quality substrate for decomposers, physically protects cellulose and other cell wall constituents, thereby reducing their degradation. A threshold level of 150 g kg–1 lignin has been suggested above which decomposition is impaired. Polyphenols include hydrolysable and condensed tannins. Insoluble condensed tannins bind to cell walls and proteins and make them physically or chemically less accessible to decomposers. Soluble condensed and hydrolysable tannins react with proteins and reduce their microbial degradation and thus nitrogen release (Mafongoya et al., 1998). A critical soluble polyphenol content of 40 g kg–1 above which nitrogen release patterns are affected has been identified (Palm et al., 2001). The relationship between polyphenol contents and nitrogen dynamics can be improved by measuring the protein-binding capacity of the polyphenols (Handayanto et al., 1994). Both lignin monomer analogues and tannins have also been shown to directly inhibit the synthesis and activity of cellulolytic enzymes (Sinsabaugh et al., 1991). Several indices of mass loss and nitrogen release from biomass have been proposed. Of these the (lignin + polyphenol):N ratio has been suggested as the most robust for materials commonly used in agroforestry, although its application still suffers from methodological difficulties in the polyphenol measurements. Such indices are not constant for plant species, because nitrogen, lignin and polyphenol contents vary for the same species with plant part, age and growth conditions including soil fertility and water supply (Mafongoya et al., 1998). Furthermore, there is evidence of rapid increases in tannin contents in the foliage of some trees following damage (Leng, 1997). For leguminous nodules, the ammonium and hexosamineN contents were the best predictors of nitrogen release during decomposition (Wardle and Greenfield, 1991). Like the elemental contents


137

Characteristics of Organic Resource N > 25 g kg–1

yes

no

Lignin < 150 g kg–1 Phenol < 40 g kg–1

yes Incorporate directly with annual crops

no Mix with fertilizer of high quality

Lignin < 150 g kg–1

yes

no

Mix with fertilizer or add to compost

Surface apply for erosion and water control

Fig. 6.1. ‘Researcher’s decision tree’ for optimum use of biomass as a function of its quality (reproduced with permission from Palm et al., 1997).

of biomass, the concentrations of organic constituents change during decomposition; for example, polyphenol contents of different litter types have been shown to decrease relatively rapidly (Pereira et al., 1998). A decision tree for the optimum use of biomass as a function of nitrogen, lignin and polyphenol contents has been proposed, where high-quality materials are directly incorporated in the soil as a nutrient source for annual crops, while materials of intermediate quality are applied together with fertilizer or high-quality biomass or are composted, and slowly decomposing, nutrient-poor materials are applied to the soil surface for erosion control (Fig. 6.1). In the absence of chemical analyses, the quality characteristics can also be estimated from simple field tests (Fig. 6.2). Leaf colour

yellow

green Leaves fibrous (do not crush) Highly astringent taste

no Incorporate directly with annual crops

yes Mix with fertilizer of high quality

Leaves crush to powder when dry

yes

no

Mix with fertilizer or add to compost

Surface apply for erosion and water control

Fig. 6.2. ‘Farmer’s decision tree’ for optimum use of biomass as a function of its quality (reproduced with permission from Giller, 2000).

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A further potentially relevant measure of biomass quality is the soluble carbon content, which may be important for initial effects on microbial growth, and thus nutrient mineralization and immobilization. Its amount in plant materials is usually 0.1 µm in the topsoil, thereby reducing the amount of plant-available water (Chauvel et al., 1991). Physical soil degradation caused by mechanized land clearing has also been described for an Ultisol in the Peruvian Amazon (Alegre et al., 1986) and an Alfisol in West Africa (Hulugalle, 1994) and contributed to yield losses in the second rotation of an oil palm (Elaeis guineensis) plantation on a ferralitic soil in the Côte d’Ivoire (Caliman et al., 1987). Potential of trees to improve the structure of agricultural soils Soil structure is stabilized by the presence of binding materials such as clay, if flocculated by polyvalent cations such as calcium or aluminium, lime, and oxides and oxihydroxides of iron and aluminium (Payne, 1988). The action of these binding materials is not influenced by the introduction of trees to agricultural systems and so will not be further discussed here (see also Section 10.3). However, the potential of trees to contribute to the maintenance and rehabilitation of soil structural characteristics in other ways has been well established. Soil physical characteristics have often been improved in hedgerow intercropping plots compared with control plots with annual crops (Rao et al., 1998). The form of these improvements has included better soil aggregation, lower bulk density, lower resistance to penetration, reduced surface sealing, improved soil porosity and consequently higher water infiltration and water-holding capacity. Similar positive effects have been found under planted fallows. The integration of trees in agricultural systems affects soil structure through the following mechanisms. •

Trees may increase the quantity of litter or mulch on the soil surface, which prevent rain drops hitting and breaking down soil aggregates, which leads to clogging of infiltration paths. Reduced overheating and evaporation from the soil surface are added advantages of a litter layer.

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•

193

In the Nigerian rainforest zone, negative effects of annual cropping on soil structure were much reduced when the soil was mulched, as this avoided crusting and maintained a higher earthworm activity (Juo and Lal, 1977; Juo et al., 1995; see also Chapter 16). Trees can increase soil organic matter levels and microbial activity through increased inputs of above- and below-ground biomass, reduced soil temperature and reduced erosion (see Chapters 4 and 17). Soil organic matter has a stabilizing effect on soil aggregates, and the loss of humus associated with cultivation is usually accompanied by structural degradation. Addition of easily decomposable organic residues to soils, such as certain tree prunings, leads to the synthesis of polysaccharides and other organic compounds that stabilize soil structure (Stevenson and Cole, 1999). Hyphae of saprophytic and mycorrhizal fungi can bind mineral and organic particles together into stable aggregates (Tisdall et al., 1997), and part of the stabilizing effect of plant roots on the soil structure is due to their association with mycorrhizas (Miller and Jastrow, 1990). Microbial polysaccharides produced in the rhizosphere could also contribute to this effect (Angers and Caron, 1998). The introduction of trees to agricultural systems will influence the quantity and types of roots present in the soil, which may affect soil structure. Plant root systems stabilize soil structure by enmeshing aggregates, releasing binding materials (mucilage) into the rhizosphere (Morel et al., 1991) and increasing soil microbial activity. As a result, root length or mass may correlate significantly with soil aggregation (Miller and Jastrow, 1990). When roots grow through the soil, they enlarge existing pores and create new ones, but their radial pressure may, at the same time, compress the soil around these pores. Water uptake by roots also affects the formation and fragmentation of aggregates by influencing wetting–drying cycles in the surrounding soil (Angers and Caron, 1998). The relatively large, continuous macropores resulting from root growth can play an important role in soil aeration and rapid macropore flow of water through the soil (see Chapter 11). In addition, old root channels are often used by new roots, which thereby avoid the mechanical resistance of the soil matrix and may profit from the more favourable chemical environment provided by root debris (van Noordwijk et al., 1991a). Plants differ in their ability to penetrate compact soil, and the increased soil porosity caused by plants with strong root systems, including certain tree species, could improve the root development of associated or subsequent crops with weaker root systems and thereby their access to additional soil water and nutrients. Like other macropores, such biopores could be especially important in facilitating root penetration when the soil matrix is too dry or its strength too high to allow rapid

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root growth. This can be essential for the establishment of seedlings (Cornish, 1993). This biological drilling by plant roots (and soil fauna) is most important at sites with compact subsoil horizons, and trees and woody shrubs are especially effective because of their perennial nature and their known ability to penetrate very hard horizons. However, in their critical review of the subject, Cresswell and Kirkegaard (1995) point out that there is not always a clear causal relationship between crop yield increases in rotations with deep-rooting species (both herbaceous and woody) and the creation of macropores in compact subsoil. To demonstrate biological drilling as the reason for yield improvements under these conditions, it would be necessary to separate effects of improved subsoil structure from those of altered soil chemical properties or the carry-over of root diseases. In future studies of the topic, particular attention should be given to the hydrological conditions of roots growing in existing macropores as influenced by possibly incomplete root–soil contact (e.g. small crop roots growing in wide tree root channels) as well as the possible concentration of pathogenic microorganisms and nematodes in such biopores (Cresswell and Kirkegaard, 1995). Methods of studying interactions between roots and soil structure have been reviewed by van Noordwijk et al. (1993b). Through the quantity and quality of root and shoot litter, microclimatic effects and avoidance of soil tillage trees may influence the abundance, composition and activity of the soil fauna, such as earthworms, termites and ants, which in turn may affect soil structure through their burrowing activity. Faunal effects on soil structure are further discussed in Chapter 16.

These strongly interdependent mechanisms deserve the attention of agroforestry researchers because they provide a basis for the use of trees in the management of soil structure. A quantitative understanding of these mechanisms is necessary for the identification of criteria for agroforestry tree species that are efficient in the maintenance and regeneration of soil structure. On a sodic soil in India, improved soil physical conditions under Prosopis juliflora compared to Acacia nilotica was explained by the higher litterfall and, more importantly, the larger root system of P. juliflora (Garg and Jain, 1992). A comparison of nine leguminous tree species and a spontaneous control in central Côte d’Ivoire provided indications that fallow species with high root mass are more efficient in the rehabilitation of compacted soils than species with lower root mass (Schroth et al., 1996). Afforestation of grassland in the Philippines with Acacia auriculiformis improved topsoil physical characteristics and water infiltration, whereas Pinus kesiya did not have this favourable effect, apparently because of the high faunal activity and well-developed root system of A. auriculiformis as

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opposed to the dense fungal mycelia in the soil associated with P. kesiya (Ohta, 1990). More mechanistic studies relating tree characteristics to their effect on soil structure are clearly desirable.

10.2 Methods for Soil Bulk Density (W.G. Teixeira, B. Huwe) Bulk density is a simple measure of soil structure. It is defined as the ratio of the mass of an oven-dry soil sample (dried at 105°C to constant weight) to its bulk volume. It is a temporally and spatially variable soil property that can be used as an indicator of changes in soil structure caused by agricultural management, root growth and activity of soil flora and fauna. In this section, only field sampling procedures will be discussed. For detailed laboratory procedures see Blake and Hartge (1986) and McIntyre and Loveday (1974). Examples of agroforestry studies in which different plant species and management practices caused significant changes of bulk densities in topsoil and subsoil can be found in Torquebiau and Kwesiga (1996), Mapa and Gunasena (1995) and Huxley et al. (1994). A reduction of soil bulk density is frequently interpreted as an improvement of soil physical properties, which is in most cases appropriate. Nevertheless, a reduction in bulk density (and increase in porosity) can sometimes be disadvantageous, for example when increased water infiltration and percolation cause increased nutrient leaching. Also, the plant-available water capacity of coarse-textured soils may be lower at low than at high bulk density, and can then be increased by compacting the soil (Archer and Smith, 1972).

Core method The soil sample is collected by driving a steel cylinder into the soil to the desired depth. The cylinder is then carefully removed to obtain an exact volumetric sample. Sampling is normally carried out by hammering or jacking. Especially for soils with large silt and clay content it is important to use appropriate samplers to avoid compaction of the sample (McIntyre and Loveday, 1974). The core diameter should be at least 7.5 cm and preferably 10 cm if accuracy is critical, and the core length should ideally be equal to the diameter (McIntyre, 1974). Sampling should be carried out at intermediate soil moisture conditions because sampling during very wet conditions may cause compression of the sample, and sampling during very dry conditions may result in shattering of the cores. In swelling soils,

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bulk density depends on soil moisture, and bulk density data should thus either be accompanied by moisture data at the time of sampling or, preferably, be determined at a specified matric suction. A suitable moisture content for measuring bulk density in such situations is when lateral swelling has reached a maximum and all cracks are closed (McIntyre, 1974). If the aim of the sampling is only the measurement of bulk density, it is not necessary to keep the collected samples undisturbed or to avoid soil moisture loss.

Clod method In situations where the sampling of cores is not practicable, as in gravelly soils, bulk density can be measured using soil clods. For such soils it is generally better to make measurements on a large clod with small disturbance than on several small cores that may have undergone serious disturbance during collection (McIntyre, 1974). The two available techniques for measuring the bulk density of soil clods, paraffin wax coating and kerosene saturation, are only suitable for relatively stable clods. These techniques apply Archimedes’ principle to determine clod volume by weighing the clod in air and in a liquid of known density. Particularly for expansive soils, this method can lead to an underestimation of bulk density due to expansion of the clod when restraining forces caused by the overlying soil are removed (Blake and Hartge, 1986). Expansive soils must therefore be equilibrated with a standard matric suction for comparison of their densities (e.g. 100 hPa; McIntyre and Loveday, 1974).

Other methods Field methods suitable for gravelly soils involve the excavation of a quantity of soil for drying and weighing and the determination of the volume by filling the hole with sand or placing a rubber balloon in it which can then be filled with water (Blake and Hartge, 1986; Liu and Evett, 1990). Nuclear techniques are also available but their use is common only in engineering applications.

10.3 Methods for Aggregate Stability (M. Grimaldi) The aggregate stability of soil affects infiltration of water and susceptibility to water erosion, crust formation, hardsetting and compaction (Angers and Mehuys, 1993). A large number of methods for evaluating the aggregate stability of soils have been proposed since the pioneering work

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of Yoder (1936). These testify, on the one hand, to the importance of this soil characteristic, and demonstrate, on the other hand, the difficulty of defining a universally applicable method. This difficulty arises because of the diversity of factors and mechanisms of disaggregation which act on different organizational levels of soil structure, and which should be reflected by the measurement. The principal mechanisms of disaggregation are slaking of aggregates by the compression of entrapped air, disaggregation by differential swelling of clays which provokes a microfissuration of the aggregates, disaggregation by the impact of rain drops and physicochemical dispersion. The stability of aggregates depends on their mineral and organic constituents (Emerson and Greenland, 1990). Three types of constituents are particularly important: the percentage of exchangeable sodium (Shainberg, 1992), the oxides and oxihydroxides of iron and aluminium (Le Bissonnais and Singer, 1993), and soil organic matter (Chenu, 1989; Haynes and Swift, 1990). Of these, it is mainly the dynamics and distribution of soil organic matter that can be influenced by trees (see Section 10.1). A method of measuring aggregate stability that has recently been proposed by Le Bissonais (1996) integrates key aspects of the older methods while being adapted to a wide range of different soil types. It consists of three main treatments which simulate the three principal mechanisms of aggregate fragmentation: • • •

treatment 1: disaggregation by slaking, provoked by rapid immersion in water; treatment 2: disaggregation by microfissuration, provoked by slow capillary wetting; treatment 3: mechanical disaggregation, provoked by shaking in water after slow capillary wetting.

The initial physical conditions of the samples, and especially their moisture and the size of the aggregates, have an important influence on the results of the treatments. These conditions are therefore standardized to allow the comparison between different samples, e.g. for different pedological horizons, climatic conditions or cropping systems. The specific soil characteristics such as their water content at the time of sampling and the plot history are taken into consideration when interpreting the results. The soil samples are sieved to obtain aggregates of a size between 3 and 5 mm, followed by air-drying or equilibration at a certain matric potential. For treatment 1, the aggregates are immersed for 10 min in distilled water. This treatment can also be used as a qualitative, simple and rapid field test. For treatment 2, the aggregates are placed on a filter paper that is wetted with distilled water, and for treatment 3 they are wetted with ethanol and then manually shaken in distilled water. Wetting in ethanol

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reduces the slaking of the aggregates by reducing the speed of the water penetration and the swelling of the clay minerals. For each treatment, and thus for each of the three mechanisms of disaggregation, the distribution of the aggregates in seven size classes is determined (>2 mm, 2–1 mm, 1–0.5 mm, 0.5–0.2 mm, 0.2–0.1 mm, 0.1–0.05 mm and 0.2 µm) Origin of the samples Primary forest Three-year-old pasture (manual deforestation) Pasture abandoned for 5 years

Mechanical compression (103 kPa) in the laboratory Mechanical deforestation and Desmodium ovalifolium cover crop Mechanical deforestation and Pueraria phaseoloides cover crop

Depth (cm)

Volume of micropores (0.2 µm > deq)

Water desorption

Mercury porosimetry

Water desorption

Mercury porosimetry

5–10

0.122

0.078

0.338

0.241

5–10 0–5 5–10 0–20

0.074 0.046 0.088 0.066

0.041 0.025 0.042 0.055

0.349 0.236 0.253 0.277

0.250 0.196 0.204 0.212

–

0.062

0.027

0.334

0.234

10–15

0.046

0.025

0.371

0.265

10–15

0.087

0.109

0.359

0.260

205

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content. This role of organic matter is particularly important in sandy soils where clay minerals and iron and aluminium oxides contribute little to the stabilization of the soil structure. Such soils are widespread in West African savannas. Based on the analysis of 495 soil samples from this region, a critical level of soil organic matter, S, was developed, from which the structural stability of the soil and its sensitivity to erosion can be deduced (Pieri, 1989): S = soil organic matter (%) / (clay + silt) (%) × 100

(10.4)

A value of S ∆LFR

P = ∆LFR + ∆DFR M = ∆DFR D=0 P = ∆LFR M=0 D = –∆DFR

P = ∆LFR + ∆DFR P=0 M = ∆DFR M = –∆LFR D=0 D = –∆LFR – ∆DFR P=0 M = –∆LFR D = –∆LFR – ∆DFR

∆LFR > ∆DFR

with permission from Fairley and Alexander (1985).

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the max–min and the balancing transfers method would indicate zero production (Kurz and Kimmins, 1987), although this situation could in theory be the result of a perfect equilibrium between root production, mortality and decomposition. Root production could also be underestimated if sampling does not coincide with seasonal peaks and troughs in root mass (Publicover and Vogt, 1993). In comparisons of agroforestry with annual cropping systems, this bias is particularly problematic because root growth and mortality are likely to be more separated in time for annual plants than for trees, and root turnover could thus be more strongly underestimated in the agroforestry plots than in the agricultural control plots (Schroth and Zech, 1995b). The third and, theoretically, the most satisfactory approach to estimate root production is the compartment flow method (Santantonio and Grace, 1987; Publicover and Vogt, 1993), which considers all three terms of Eq. 12.3. With this method, mortality (M) is calculated from changes in DFR and a decomposition constant k, assuming an exponential decomposition of the dead root mass. Production and decomposition are then calculated by transforming Eqs 12.1 and 12.2 above (Publicover and Vogt, 1993): M = kt[DFRt2–DFRt1e(-kt)] / [1 – e(-kt)]

(12.4)

P = LFRt2 – LFRt1 + M

(12.5)

D = DFRt1 – DFRt2 + M

(12.6)

The decomposition constant k is related to the disappearance rate DR (i.e. the fraction of dead roots that disappears during a certain time interval such as a month) by: k = – ln (1 – DR)

(12.7)

Equation 12.4 can be modified if root decomposition is known to differ from the simple exponential model. In theory, the compartment flow method allows accurate estimation of root production even when root growth, mortality and decomposition occur simultaneously. The principal problem with this approach is the need for an accurate decomposition rate for fine roots, which is difficult to obtain (see Section 6.3). Under agroforestry conditions, the decomposition rate would depend on the relative contribution of tree and crop roots, including its seasonal fluctuations, to the dead root pool, as these roots are likely to differ in their decomposition dynamics. Unfortunately, the method is relatively sensitive to errors in the decomposition rate. It may, therefore, be useful to provide production estimates for a range of decomposition rates (Schroth and Zech, 1995b). The method is also sensitive to misidentification of live and dead roots, which are often not easy to distinguish, especially for tree species with dark roots, and to loss of dead roots when processing the

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samples. However, modelling results indicate that the production estimates remain more accurate than those of the other two calculation methods unless the processing error becomes quite large (Publicover and Vogt, 1993). Whenever possible, the compartment flow method should be used for calculating root production from sequential soil core data, possibly in combination with the balancing transfers method for comparison. As indicated above, the max–min and the balancing transfers methods have an inherent tendency to underestimate root production because they ignore the simultaneous occurrence of root production, mortality and decomposition. However, there is another source of error with these methods which may lead to an overestimation of root production. This error arises from the summation of random increases in root mass between sampling periods. It increases with increasing sampling frequency and is especially severe for sites where there are no pronounced seasonal changes in live and dead root mass. To avoid this error, it is recommended to use only statistically significant changes in root mass for production calculations with these two methods (Vogt et al., 1998). This is a serious limitation because it requires the separate processing of many replicate root samples which could otherwise be pooled and subsampled to increase the precision of the measurement that can be achieved per unit processing time (Schroth and Kolbe, 1994). In some cases this problem can be circumvented because it is known, or at least it is very likely, that roots have turned over, even if differences between sampling dates are not significant at a given level of replication. This would be the case when tree roots show pronounced seasonal fluctuations in accord with weather or management actions such as pruning, in systems with annual crops whose roots necessarily die after the harvest (Schroth and Zech, 1995b), or when additional observations (e.g. with rhizotrons, see below) indicate root turnover between sampling dates (Vogt et al., 1998). However, where such evidence is not available and root data show no pronounced seasonal patterns, as for perennial crops or trees in climates without pronounced seasons, the max–min and the balancing transfers methods should not be used for production calculations. Modelling results indicate that the compartment flow method is not prone to this source of error (Publicover and Vogt, 1993), and it is therefore not necessary to restrict root production calculations to significant changes in live and dead root mass. The compartment flow method is ideally combined with the pooling–subsampling method to reduce processing time (Schroth and Kolbe, 1994). As indicated before, however, further research on root decomposition rates for different trees and crops as influenced by soil and climatic conditions and their mathematical description is required to make full use of the potential of this method.

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Sequential excavation As soil coring is only suitable for sampling fine roots, other methods are required when coarse roots are also of interest, such as in studies of total carbon accumulation or progressive soil exploration by coarse lateral or vertical roots. Dynamics of coarse roots can be studied by sequential excavation of trees of different ages, which may be combined with soil coring for quantification of fine roots. Sequential excavation was used, for example, by Yocum (1937) for coarse and fine roots in his studies of apple tree root development as influenced by associated crops and soil management practices. After suitable calibration, allometric techniques or simply the root–shoot ratio can be used for estimating coarse root biomass of trees (van Noordwijk et al., 1996). These techniques have rarely been used in agroforestry because of the considerable work involved and the focus on fine roots in most studies. They could, however, become more relevant with increasing attention given to the potential of trees to increase carbon sequestration in agricultural systems.

Root ingrowth method This method is best suited for comparative studies of root growth for different time periods, sites or experimental treatments, although the method has also been used for estimating root production in tropical forests (Jordan and Escalante, 1980; Kangas, 1992) and shaded tree crop plantations (Muñoz and Beer, 2001). A certain soil volume (usually a core, but sometimes a soil block) is removed and replaced by root-free soil in a mesh bag, so that root ingrowth during the incubation period into a defined soil volume can be determined. The mesh bag can be made of nylon or metal netting with an approximately 5 mm mesh (Steen, 1991; Muñoz and Beer, 2001). The soil in the ingrowth core is either root-free soil from the same site, or a standard substrate such as sand or vermiculite. If soil from the site is used, live and dead roots are removed as far as possible by dry sieving. The use of a polyethylene sheet, charged with static electricity by rubbing, can help in the removal of fine root fragments (Muñoz and Beer, 2001). These procedures will not produce a completely root-free soil, so root residues should be extracted from a subsample for correction of the final data of root ingrowth (Persson, 1990). Incubation periods of 1–12 months have been used in humid tropical (Cuevas and Medina, 1988; Raich et al., 1994; Muñoz and Beer, 2001) and Mediterranean climates (Fabiao et al., 1985). Depending on the research objective, either all cores are collected after a fixed incubation period, or sets of cores are collected after increasingly long incubation periods (e.g. 3, 6, 9 and 12 months) to study the progress of root ingrowth into the

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cores. In the absence of site- and species-specific experience with the method, the latter approach is probably preferable. With the ingrowth technique, Fabião et al. (1985) studied the seasonal dynamics of root growth in Eucalyptus globulus plantations in Portugal, and Muñoz and Beer (2001) studied root growth dynamics and turnover in shaded cocoa plantations in Costa Rica. The ingrowth method is well suited for studying the effect of experimental soil manipulations on root growth. For example, Cuevas and Medina (1988) and Raich et al. (1994) demonstrated limitation by certain nutrients for forests in Amazonia and Hawaii, respectively, through increased root ingrowth into cores with standard growth media to which these nutrients had been added. With a similar design, McGrath et al. (2001) could not clearly detect phosphorus deficiency in an Amazonian agroforestry system despite very low soil phosphorus availability. Steen and Hakansson (1987) demonstrated reduced root growth of annual crops into cores with increasing soil compaction. A disadvantage of the method is that, unlike sequential soil coring, it does not provide information on root distribution in the soil. If this information is required, the method has to be combined with another method, such as soil coring (Muñoz and Beer, 2001) or the profile wall method. If quantitative growth data are needed, a potential problem is the difficulty of creating conditions in the core which are similar to those in the surrounding soil with respect to factors such as soil structure (and thus water content), nutrient mineralization and the presence of competing roots. This is much easier to achieve in a recently tilled topsoil of an agricultural field than in a plot under permanent vegetation. In agroforestry studies with several contrasting plant species, such as annual crops and trees, artificial growth conditions in the cores could affect plant species differently, thereby introducing bias. If ingrowth cores are used for measuring root production, information on live roots, dead roots and root decomposition should be taken into consideration, as outlined above for sequential soil coring (Steen, 1991). Several authors found reasonable agreement of production values obtained with the ingrowth and sequential soil coring methods (Persson, 1983, for a boreal pine forest; Hansson and Andrén, 1986, for a perennial grass ley; Symbula and Day, 1988, for a swamp forest), whereas others found substantial differences in fine root production (Neill, 1992, for a prairie marsh; Makkonen and Helmisaari, 1999, for a boreal pine forest). Until research in tropical agroforestry systems provides evidence that (or under what conditions) root production estimates from ingrowth cores are reliable, it is recommended that this technique is used for comparative purposes or in combination with other methods for measuring root production.

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Minirhizotrons Minirhizotrons are a non-destructive method for the in situ observation and quantification of roots in the soil. They consist of a transparent access tube, such as a glass or acrylic tube, or a hole in the soil, through which images of roots growing at the interface with the soil can be obtained with a fibre optic probe, miniaturized video camera, or simply a mirror and a camera with a macro-lens (Mackie-Dawson and Atkinson, 1991; van Noordwijk et al., 1996). Johnson et al. (2001) provide a recent review of the technique. Minirhizotrons have been used for collecting data on root length distribution, root dynamics (growth, senescence and decomposition), root properties (branching patterns and suberization) and interactions of roots with other organisms (herbivory, parasitism and symbioses such as root nodules formed with rhizobia). Due to their non-destructive nature, they allow roots to be monitored through different seasons, wetting–drying cycles, phenological cycles of a plant, or in response to management interventions such as shoot pruning or fertilization (Ball-Coelho et al., 1992; Kätterer et al., 1995; van Noordwijk et al., 1996; Price and Hendrick, 1998; Vogt et al., 1998). If roots of different species can be distinguished, the technique allows comparative studies of root dynamics of associated trees and crops or pastures in agroforestry associations, which may give important information on below-ground interactions between species (Campbell et al., 1994). Often, however, it is not possible to distinguish roots of associated species (Hendrick and Pregitzer, 1996). Several software packages are available for analysing root data from minirhizotron observations (Vogt et al., 1998). When working with minirhizotrons, an important consideration is how to avoid artefacts due to altered growth conditions of the roots at the interface between minirhizotron and soil. In sandy, uniform, well-packed soil, the presence of rigid observation tubes does not seem to influence root growth (Vos and Groenwold, 1987; Ephrath et al., 1999), although preferential root growth at the soil–tube interface may occur in finertextured soils. In compact soil, this may lead to deeper root growth along the tubes than in the bulk soil (Vos and Groenwold, 1987). To reduce the problem of gap formation at the interface between rigid observation tubes and soil, several types of inflatable minirhizotron tubes have been tested. Gijsman et al. (1991) described a system consisting of a rubber tube, actually a modified motorcycle inner tube, which exerts a small pressure on the surrounding soil when inflated, thereby avoiding gaps at the interface, and is removed when observations are made. Advantages include the better visibility of the roots, which are not obscured by an imperfectly transparent tube, and the elasticity of the tube, which is important in swelling and shrinking soils (Gijsman et al., 1991; van Noordwijk et al., 1996).

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Another factor to be considered is the installation angle of the tubes. Root growth along the side of minirhizotrons has been observed with vertically installed, rigid tubes. Sometimes, but not always, this can be avoided by installing the tubes at an angle of 45° to the soil surface (MackieDawson and Atkinson, 1991). However, in studies of deep root systems, as with many trees, installation angles other than vertical are impractical (Ephrath et al., 1999). In a study with Acacia saligna and wheat in a sandy, disturbed soil, Ephrath et al. (1999) found no difference in root length density measured with tubes placed vertically or at an angle of 45° in the soil. As discussed above, larger effects of the installation angle would be expected in more clayey soils. In studies with shallow-rooted crops, horizontal installation of minirhizotrons may be advantageous because it leads to more images per soil depth, thereby reducing variability of mean values (Dubach and Russelle, 1995) and allowing the analysis of spatial patterns. Johnson et al. (2001) provide a detailed discussion of installation methods for rhizotron tubes and recommend a waiting period of 6–12 months between installation and data collection in order to avoid artefacts caused by soil disturbance. Light can negatively affect root growth, even at low intensities, and so its entrance into the tubes needs to be avoided (Caldwell and Virginia, 1989). Also, temperature effects should be avoided by insulating exposed ends of rhizotron tubes above the soil surface (Samson and Sinclair, 1994). The minirhizotron technique is well suited for comparative measurements of root dynamics in different seasons or experimental treatments. Repeated observations of the same root can provide information on root longevity and turnover. Johnson et al. (2001) recommended sampling intervals of 2 weeks or less in such studies in order to reduce underestimates of root turnover, which occur when roots appear and disappear between two sampling events. Difficulties have been encountered when attempting to transform minirhizotron readings into quantitative values of root length or root mass per unit bulk soil. Calibrations of minirhizotron observations against soil coring data produced relationships that varied with plant species and phase of crop development (Ball-Coelho et al., 1992; Merrill and Upchurch, 1994; Samson and Sinclair, 1994). Furthermore, for both herbaceous crops and trees, correlations between these methods tend to be poorest near the soil surface, where root length is often underestimated by the rhizotron technique (Ball-Coelho et al., 1992; Samson and Sinclair, 1994; Ephrath et al., 1999). This is problematic, because most roots are usually concentrated here. A theoretical calibration approach for estimating root length from minirhizotron data also produced contradictory results (Merrill and Upchurch, 1994). More encouraging results were recently obtained for estimating root mass of maize and two temperate tree species from minirhizotron observations of root surface area (Jose et al., 2001).

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Despite these difficulties, the minirhizotron technique can be useful in the estimation of root production if the turnover values obtained from repeated observations of roots at different depths are combined with absolute root mass data as obtained, for example, with soil coring at one or several points in time. Studies that have used this approach have been reviewed by Hendrick and Pregitzer (1996).

Nutrient fluxes associated with root turnover Nutrient fluxes through fine root systems of trees are often calculated by multiplying estimates of root production with nutrient concentrations in root samples, assuming that there is no retranslocation of nutrients from senescing roots (Burke and Raynal, 1994; Lehmann and Zech, 1998; Muñoz and Beer, 2001). Available evidence on this topic is contradictory. A study by Nambiar (1987) suggested little translocation of nitrogen, phosphorus, potassium and magnesium from senescing fine roots of Pinus radiata, but other studies have indicated significant retranslocation of nitrogen and phosphorus from senescing roots of northern conifers (Meier et al., 1985; Ferrier and Alexander, 1991). A recent analysis of a large data set of nutrient concentrations in live and dead roots found no evidence for retranslocation of nitrogen, but indicated an average retranslocation of about 30% of phosphorus and potassium from senescing fine roots. For magnesium the average tissue concentration decreased by 25% between live and dead fine roots, but the difference was not significant (Gordon and Jackson, 2000). These results suggest that the turnover of phosphorus, potassium and perhaps magnesium in fine roots may be overestimated if calculated as indicated above, although estimates of nitrogen turnover in fine roots may in many cases be reliable. Further research is needed to confirm this. If roots of annual crops die after harvest or tree roots are sloughed off by tillage, retranslocation of nutrients obviously does not occur.

Chapter 13 Biological Nitrogen Fixation K.E. GILLER Plant Production Systems, Department of Plant Sciences, Wageningen University, PO Box 430, 6700 AK Wageningen, The Netherlands

13.1 Synopsis Trees can increase nutrient inputs and reduce nutrient losses through a variety of processes. Biological nitrogen fixation is prominent among these as it is the only truly renewable source of nutrients in agroforestry or agriculture as a whole. Many of the trees used in agroforestry are legumes as the ability to fix nitrogen allows them to grow rapidly in nitrogendepleted soils. Thus one of the principal roles of nitrogen-fixing trees in agroforestry is to improve soil fertility, athough many legume trees have multiple uses for fodder, fuelwood, fruits and timber. Legume trees are thus found in almost all types of agroforestry systems: as shade trees in perennial crops, in improved fallows, in hedgerow intercropping systems, as erosion barriers, live fences, isolated trees in parklands and fodder trees. Not all legumes can nodulate and fix nitrogen, and the taxonomy of the Leguminosae is a good guide to which legume trees are nitrogen fixers. Nitrogen-fixing symbioses are also formed between non-legume trees and actinomycetes (Frankia spp.), which are termed actinorhizal symbioses (Giller, 2001). The most important of these are the Casuarina spp. from Australasia, which are used for soil stabilization, as windbreaks and for poles and fuelwood throughout the tropics. The actinorhizal symbioses have been less studied in agroforestry than leguminous trees, but the methods outlined here can equally well be applied to the study of either type of symbiosis. A full list of nitrogen-fixing trees also includes larger species of cycads, which form symbioses in their leaf axils with blue-green algae (Cyanobacteria), but these are of no importance in agroforestry. © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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Nodulation and nitrogen fixation in the Leguminosae The Leguminosae are a large and diverse family of flowering plants composed of some 18,000 species, ranging from huge, long-lived forest trees to tiny, annual, herbaceous forbs. All legumes have nitrogen-rich tissues, irrespective of whether they can fix nitrogen, and it is now thought that the nitrogen fixation ability evolved to satisfy the demand for nitrogen, rather than nitrogen fixation resulting in the high tissue concentrations of nitrogen, which had often been assumed earlier (see McKey, 1994; Giller, 1997). Those legumes, which are not able to fix nitrogen, also have a large tissue nitrogen demand, which they satisfy either by competing actively for soil nitrogen or by growing more slowly and conserving their nitrogen within the plant. A further major consequence of the large tissue concentrations of nitrogen in the Leguminosae is the wide variety of secondary metabolites which are produced to protect legume tissues from predation (Waterman, 1994). Among these secondary metabolites the polyphenols, and particularly the condensed tannins, play a major role in regulating nitrogen release from legume tissues in decomposition (see Sections 6.1 and 6.4) and during ruminant digestion (Giller, 1997). The Leguminosae are divided into three subfamilies, the Caesalpinioideae, the Mimosoideae and the Papilionoideae, and these subfamilies provide a useful guide as to whether the legumes are able to nodulate and fix nitrogen. The Caesalpinioideae, of which only 23% form nodules (de Faria et al., 1989; Sprent, 2001), are considered to be the most primitive group. Within the Caesalpinioideae, nodulation is restricted to a few tribes, the most notable of which is Cassieae, which appear to form a bridging group between the non-nodulating and nodulating legumes. Within this tribe all members of genera Cassia (sensu stricto) and Senna are unable to nodulate. The genus Senna includes several important agroforestry species, such as S. siamea and S. spectabilis, which have been widely tested in hedgerow intercropping and were earlier assigned to the genus Cassia (Irwin and Barneby, 1981). Closely related to the genera Cassia and Senna is the genus Chamaecrista, in which all the species examined nodulate and fix nitrogen. The range of nodule morphology within the genus Chamaecrista is striking, with some species having rather primitive nodule structures, in which the bacteria are maintained in persistent infection threads within the nodule cells, whereas in other species fully differentiated bacteroids are released within peribacteroid membranes in the host cell cytoplasm (Naisbitt et al., 1992). Persistent infection threads are also found in the only non-legume tree genus, Parasponia (Ulmaceae), which can form nodules and fix nitrogen with rhizobia (Trinick and Hadobas, 1988; Becking, 1992). Almost all legumes in the other two subfamilies, the Mimosoideae (90%) and the Papilionoideae (97%), can form nodules with rhizobia and fix nitrogen (de Faria et al., 1989). The exceptions in the Mimosoideae appear

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to be cases where loss of nodulation and nitrogen-fixation ability has occurred in species which are found in very arid environments (e.g. some Acacia spp.), presumably due to the sensitivity of nitrogen fixation to drought (Odee and Sprent, 1992; Sprent, 1994). Members of the Papilionoideae which do not nodulate are largely those found in primitive tribes. Useful reference texts with information on nodulation and nitrogen fixation in legumes are Allen and Allen (1981) and Sprent (2001).

Other sources of nitrogen fixation in agroforestry systems Apart from inputs from nitrogen-fixing trees, herbaceous cover crops/ green manure legumes and grain legumes are often important components of agroforestry systems and further information can be found in Giller (2001). The discovery of endophytic nitrogen-fixing bacteria in the tissues (particularly the vascular tissues) of a range of non-legume species, including oil palm (Elaeis guineensis), has led to claims that such bacteria may give substantial inputs of nitrogen. Direct evidence for substantial nitrogen inputs (~20 kg N ha–1 year–1) is strongest in sugar cane (Saccharum officinarum; e.g. Boddey, 1995), though the evidence is still controversial. Nitrogen inputs from heterotrophic, free-living nitrogen-fixing bacteria in soil are generally considered to be very limited and not of significance except over long time scales (Giller and Day, 1985; Vanderleyden, 1997; Giller, 2001).

Environmental limitations to nitrogen fixation All environmental limitations that adversely affect plant growth and vigour also decrease amounts of nitrogen fixation in legumes, although the symbiosis is sometimes more sensitive to such constraints than other aspects of plant growth (Giller, 2001). Nitrogen fixation is sensitive to nutrient deficiencies, in particular phosphorus deficiency, which may restrict nodule formation if acute. Molybdenum deficiency influences nitrogen fixation directly as molybdenum is a component of the nitrogenase enzyme. Nitrogen fixation is thought to be more sensitive to drought stress than other processes, such as photosynthesis, although the evidence is somewhat equivocal (Sprent, 1984). A further interesting feature of the legume–rhizobium symbiosis is that the sensitivity to stress may be expressed through the bacteria, the legume host or the formation of the symbiosis itself. Legumes are invariably more sensitive to salinity than are rhizobia (Sprent, 1984), but rhizobia are more sensitive than their hosts to heavy metal pollution (Giller et al., 1998). The infection process itself

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appears to be particularly sensitive to calcium deficiency (Giller, 2001). Large differences in sensitivity to stresses such as aluminium toxicity in soil are found among rhizobial strains and among legume hosts.

Effects of management on nodulation and nitrogen fixation Legume trees are often pruned severely in agroforestry systems to provide fodder or foliage for soil amendment. It is well established that defoliation causes nodule senescence (Witty and Minchin, 1988), and pruning or browsing of trees by animals causes temporary decreases in the rates of nitrogen fixation. Reestablishment of nitrogen fixation will depend on formation of new nodules, though this can sometimes be rapid as legumes often harbour latent infections in young roots, which can develop when nitrogen demand in the plant is large. However, in Erythrina poeppigiana pruning resulted in complete mortality of nodules and there was a lag of 10 weeks before active nodules were re-formed (Nygren and Ramírez, 1995). Similarly, pruning of Leucaena diversifolia resulted in drastic reduction of nodule activity (measured as acetylene reduction) for about 3 months (Snoeck, 1995). Nygren (1995) highlighted the danger that nodulation might be completely suppressed if trees are pruned too frequently. Nodule senescence associated with defoliation of trees is a mechanism by which nitrogen is made available to plants growing in close proximity, though amounts of nitrogen made available in this way are likely to be relatively small (Rao and Giller, 1993; Nygren and Ramírez, 1995). Even Table 13.1. Mechanisms by which nitrogen from nitrogen-fixing trees can be made available to other plants.a

Mechanism Below ground Root and nodule senescence and mineralization Rhizodeposition Transfer between roots by interconnected mycorrhizal hyphae Above ground Mineralization of severed or senesced plant material Consumption by grazing animals or insects and return in excreta or as carcasses Foliar leachates Transfer of ammonia to associated plants aModified

from Ledgard and Giller (1995).

Rate of transfer

Likely importance as an N source

Slow Rapid

Major Minimal

Rapid

Minimal

Slow

Major

Slow/rapid Rapid Rapid

Major Moderate Minimal


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Box 13.1. Rhizobia that nodulate legume trees. The rhizobia that nodulate legume trees are a sadly neglected area of study compared with the amount of research effort which has been directed to grain legumes. The taxonomy of rhizobia has developed rapidly since the advent of phylogenetic methods for bacterial classification based on sequence analysis of the 16S rRNA gene (Young and Haukka, 1996). The older classification methods were largely based on the legume host range of the rhizobia. Although this approach was recognized to be severely flawed early on (Wilson, 1944), the new classification reveals the true extent of such problems in that nodulation ability is closely related to the nodulation genes carried by the rhizobia, rather than their evolutionary similarity (Young and Haukka, 1996). The nodulation genes are generally carried on transmissible plasmids in fast-growing rhizobia, and in some cases on transmissible ‘symbiotic islands’ of chromosomal DNA (Sullivan and Ronson, 1998). Rhizobia far apart on the phylogenetic tree may thus carry the same nodulation genes and have a very similar host range for nodulation and nitrogen fixation. Research in the last 10 years has revealed some surprising overlaps between rhizobia which can nodulate legume trees, herbaceous and grain legumes. The best documented case is that of Rhizobium sp. NGR234, which has been shown to effectively nodulate legumes from 112 genera, including members of the three different subfamilies of the Leguminosae (Pueppke and Broughton, 1999). Other broad-host-range rhizobia that nodulate trees are Rhizobium tropici (MartínezRomero et al., 1991), which nodulates Leucaena spp. and Phaseolus vulgaris, and indeed other rhizobia species such as R. etli, which were thought to be specific in their host range, have since been shown to nodulate a wider range of species including trees (Hernandez Lucas et al., 1995). Slow-growing rhizobia are all members of a single genus, Bradyrhizobium, whereas the fast-growing rhizobia have been split into several genera. Five genera of fast-growing rhizobia are currently recognized: Rhizobium, Azorhizobium, Sinorhizobium, Mesorhizobium and Allorhizobium, although this field is changing rapidly. Some trees are reported to nodulate only with slow-growing rhizobia (Dreyfus and Dommergues, 1981) but the vast majority examined nodulate with fast-growing rhizobia. Of particular note are the rhizobia which form stem nodules; shrubby legumes of the genus Aeschynomene form stem nodules with photosynthetic slow-growing Bradyrhizobium strains, whereas Sesbania rostrata forms nodules with rhizobia classified into a separate genus, Azorhizobium caulinodans (Giller, 2001). Recent studies of rhizobia that nodulate legume trees have resulted in the description of the new species Sinorhizobium terangae and Sinorhizobium saheli from nodules of Sesbania and Acacia spp. in Senegal (de Lajudie et al., 1994) and Mesorhizobium plurifarium (de Lajudie et al., 1998), Sinorhizobium arboris and Sinorhizobium kostiense (Nick et al., 1999) from nodules of Prosopis and Acacia spp. in Sudan and Kenya. The classification has recently been substantially revised (see Young et al., 2001)

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with E. poeppigiana, which was profusely nodulated in the field, amounts of nitrogen released from complete senescence of the nodules were much less than 10 kg N ha–1 (Nygren and Ramírez, 1995). If the turnover of fine roots is included, the amounts of N recycled below ground are likely to be of considerable importance but accurate estimates are lacking. The major pathway by which nitrogen is made available for other plants is through recycling of above-ground litter either directly to the soil, or through manure of animals grazing on the trees (Table 13.1).

Requirements for inoculation The success of many of the fast-growing legume trees in agroforestry has undoubtedly been due to their ability to nodulate and fix nitrogen in soils across the tropics. Unfortunately, it is almost impossible to generalize as to which species are likely to nodulate readily in soils where they have not been grown, although there are gradations of the degree of promiscuity or specificity among agroforestry trees (Bala and Giller, 2001). Bala (1999) hypothesized that the success of species such as Leucaena leucocephala and Gliricidia sepium could have been due to either: (i) their ability to nodulate promiscuously with a wide range of rhizobial types, or (ii) the fact that, although such trees may nodulate with a narrow range of rhizobial species, these species are ubiquitous throughout the tropics. To test these hypotheses he characterized rhizobial isolates from nodules of Leucaena leucocephala, Calliandra calothyrsus, Gliricidia sepium and Sesbania sesban grown in soils from Central and South America, West, East and southern Africa and South-east Asia. Rhizobial isolates were characterized on the basis of physiological tests, host range for nodulation and molecular typing methods (including 16S rRNA partial gene sequences). Bala (1999) found that rhizobial isolates nodulating Leucaena, Calliandra and Gliricidia were diverse, belonging to a wide range of strains of the genera Rhizobium, Mesorhizobium, Sinorhizobium and Allorhizobium/Agrobacterium. This supports the first hypothesis that the tree legumes nodulated promiscuously with a wide range of rhizobia. Further, these three tree legumes nodulated with many of the same isolates although there were isolates which nodulated with only one or two of the species. Sesbania sesban was an exception in that it nodulated with a largely separate range of isolates, which fell into the Rhizobium and Mesorhizobium genera. Even more surprising was the limited range of soils from southern Africa in which S. sesban nodulated, considering that it is naturally found in this region. This clearly indicated that S. sesban requires inoculation when sown for the first time into upland soils. Details of how rhizobial inoculants can be made and applied are given in the methods manuals cited below.


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13.2 Microbiological Methods for Studying Rhizobia Methods for the isolation of rhizobia from nodules and authentication are simple and can be conducted in any laboratory which has the capacity for simple microbiology. There are a number of excellent methods manuals including Vincent (1970), Somesagaran and Hoben (1985) and SylvesterBradley and Kipe-Nolt (1988). Host-range studies provide information on the ability of individual isolates to nodulate and fix nitrogen with different legumes, and therefore can give some clues as to the phylogenetic relatedness of strains, but detailed strain characterization now relies on methods of molecular biology, which tends to restrict such activities.

13.3 Simple Methods for Determining Whether a Legume is Fixing Nitrogen In the absence of access to isotope-based methods (see below) there are some simple approaches to assessing whether trees or crops are benefiting from nitrogen fixation. •

•

The simplest and most obvious (though often overlooked) approach to determine whether a tree is fixing nitrogen is to excavate the root system and look for nodules. If nodules are found, then this is strong evidence that the trees are or have recently been fixing nitrogen. If no nodules are found this could be due to a number of reasons: that the root excavations are not complete and the nodules are only found in deeper horizons, or that the nodules readily become detached from the roots and lost during sampling, which is particularly problematic with some species (e.g. Cajanus cajan, Gliricidia sepium). The ability of a tree to grow and accumulate nitrogen in infertile or skeletal soils may be evidence of nitrogen fixation. Sanginga et al. (1986) compared nitrogen accumulation in prunings of inoculated and uninoculated Leucaena leucocephala and estimated that 224–274 kg N ha–1 came from nitrogen fixation. This approach to estimating nitrogen fixation, which is a simple and useful approach to estimating nitrogen benefits, is referred to as the nitrogen balance method. However, in many situations it is dangerous to rely on such evidence alone as tree roots might be accessing deep soil horizons richer in nitrogen (see Box 8.1 on p. 171), or there may be subsurface lateral flow of water from which the trees can scavenge nitrogen. A further development of this approach is to develop a whole nitrogen balance for the system, preferably comparing a system with nitrogen-fixing trees with a system with non-nitrogen-fixing trees. Such studies suffer from the large number of unknown processes such as gaseous losses,

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which are as difficult to estimate as nitrogen fixation. A primary advantage of agroforestry systems is that the trees are able to access nutrient sources which are not available to other plants, which in itself can cause problems in calculations of nitrogen balances if nitrogen is taken up from deep water tables.

13.4 Isotope-based Methods for Measurement of Nitrogen Fixation N-isotope-based methods are perhaps the only means available for estimating amounts of nitrogen from nitrogen fixation in trees growing in the field, though such methods may have significant problems in their application. The two main variants of these methods are those involving addition of 15N-labelled fertilizers (often called 15N-isotope dilution methods) and those that rely on the natural 15N-enrichments of soil (known as natural abundance methods). Most soils are naturally, slightly enriched with 15N with δ15N values commonly of 2–16‰ (the difference between the 15N content of the material and air expressed in parts per 1000): 15

 Rsample – Rreference   × 1000 Rreference  

δ15N (in parts per 1000 or 0/00) = 

(13.1)

where during chemical analysis R is the ratio of N2 molecules derived from the plant material, which are composed of one 15N and one 14N atom to those composed of two 14N atoms, i.e. R=

15

N+ 14 N

14

N+ 14 N

(13.2)

The basic principle of both methods is that nitrogen taken up from the soil is enriched with the heavy isotope 15N compared with nitrogen absorbed from the atmosphere. A non-nitrogen-fixing reference plant is used to estimate the 15N-enrichment of nitrogen available from the soil in both cases (Peoples et al., 1989). The major assumption (and problem) of these methods is that the 15N-enrichment of the available soil nitrogen taken up by the reference plant is the same as that taken up by the nitrogen-fixing legume. This is in fact only true when there is a perfectly uniform 15N-enrichment of plant available nitrogen in the soil with depth and time, or, if this is not the case, when the rooting depths and activity and the time course of nitrogen uptake of the reference and legume plants are identical (Witty, 1983). This assumption is rarely satisfied and therefore steps must be taken to limit the variability in 15N-enrichment of the available soil nitrogen as far as possible (Witty and Giller, 1991). The use of direct measurements of the 15N-enrichment of the available soil nitrogen


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coupled with modelling have been recommended as an alternative to using reference plants, but such approaches require extensive sampling and analysis of extractable soil nitrogen (Chalk and Ladha, 1999) and are unlikely to be widely used. Much has been written on the problems of measuring nitrogen fixation using isotope approaches (e.g. Witty, 1983; Witty and Giller, 1991; Chalk and Ladha, 1999). Problems encountered with herbaceous legumes when isotope methods are applied are magnified in studies with trees due to their much larger size and longevity (Boddey et al., 2000). The size of trees and lignification of plant parts presents additional problems of subsampling of plant tissues to give a representative sample for measurement of tissue nitrogen concentrations and 15N-abundance.

Box 13.2. Measuring nitrogen fixation under controlled conditions. There are remarkably few measurements of nitrogen fixation by nitrogen-fixing trees growing in the field. This is in no small part due to the technical problems that exist in the application of the available methods, and partly for this reason many measurements have been confined to studies under more controlled conditions with trees grown in pots. The major limitations of these pot studies is that they do not often provide the information we need to assess the role of nitrogen fixation in agroforestry systems. So what do we want to measure? Do we want to know which of the different provenances of Gliricidia sepium fix the most nitrogen after 6 weeks of growth in a glasshouse? Or do we wish to pose the question: are all rhizobial strains that belong to different species or genera equally efficient in nitrogen fixation with a particular tree species? If these are the types of questions raised, then experiments in pots are a valid approach, but we must further question what methods are required to address these questions. The principal reason for using the 15N-isotope dilution method is to distinguish between nitrogen derived from the atmosphere and nitrogen derived from the soil. If experiments are conducted in pots, the supply of nitrogen from the soil can be carefully controlled, so there is little advantage in using the 15N-isotope dilution method compared with simpler nitrogen balance methods. The major concern with such studies under controlled conditions in pots is that there is no guarantee that there is any correlation whatsoever between results found under controlled conditions in the glasshouse and the response of the same treatments in the field. This is largely due to the strong interactions between nodulation and nitrogen fixation and a wide range of environmental factors, which are described above.

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13.5 Estimating Nitrogen Fixation in Field Settings It can be argued that 15N-isotope dilution using labelled fertilizers has no role to play in estimation of nitrogen fixation by trees, for the following reasons. •

•

•

A wide variability in rooting patterns can be found even within a single provenance of a tree species, as soil chemical and physical properties exert a very strong influence on root distributions. As mentioned before it is difficult to ensure that 15N is incorporated to depth and unlikely that non-fixing plants can be identified that have matching rooting patterns. One of the major advantages of legume trees is that they can be deeprooting, but this raises obvious problems in uniform 15N-labelling of soil with depth. These are problems that also apply to measurement of nitrogen fixation in herbaceous legumes but are amplified in agroforestry systems because of the larger size and potentially deeperrooting systems of the trees. 15N-isotopes are expensive. Although it can be argued that the cost of the isotopes is relatively small compared with other costs of field experimentation and mass spectrometric analysis of samples, the wide planting arrangements of trees in agroforestry systems demand the use of large 15N-labelled microplots. There is, therefore, a strong temptation to try to cut costs by not including border rows in the 15Nlabelled microplots, but sampling the whole area to which 15N-labelled fertilizers have been applied. Obviously, this breaks the basic rules of experimentation with 15N-labelling (see, for example, Stumpe et al., 1989). Given the extensive rooting systems which have often been observed in agroforestry trees, border rows are particularly important (see Section 3.2).

This means that the 15N natural abundance method is probably the only sensible choice for measuring nitrogen fixation by trees in the field, but again problems should not be underestimated. The δ15N of soil nitrogen may vary markedly with depth, although the extent to which this is a problem is debatable (Boddey et al., 2000). Some studies have found little change of δ15N in the plant-available nitrogen with depth, but there is considerable variation between sites so this should be investigated if possible. It is unlikely that a reference tree can be identified which will have an identical pattern of root activity with depth, or uptake with time, as that of the nitrogen-fixing tree under investigation. One approach to this problem is to use a range of potential reference plants. This was recommended for measurements of nitrogen fixation in grain legumes (Boddey et al., 1990) but is especially applicable for nitrogen-fixing trees. If most of the reference plants give δ15N values close to the δ15N of the total


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Kjeldahl nitrogen of the soil on which they are growing, but one reference plant consistently gives δ15N values significantly different from the rest, then that is a reasonable indication that it is an aberrant reference value and may be discarded. Such a situation was found when measuring nitrogen fixation in Faidherbia albida in Malawi where mango (Mangifera indica) consistently gave small δ15N values (2.7–4.8‰) compared with the soils (6.4–11.5‰) and other non-fixing trees on the same sites (6.0–10.0‰) (Phombeya, 1999). The reason(s) why certain plants tend to become depleted or enriched in 15N relative to their sources of nitrogen are not fully understood but appear to be related to mycorrhizal colonization, drought and nitrogen deficiency (Högberg and Alexander, 1995; Handley et al., 1996, 1999). Högberg (1990) showed that ectomycorrhizal non-nodulating legume trees (Brachystegia and Julbernardia spp.) in miombo woodland of Tanzania were more depleted in 15N than arbuscular mycorrhizal-nodulating or non-nodulating legume trees. Subsequent studies in Cameroon did not confirm that these differences were due to the type of mycorrhizal infection (Högberg and Alexander, 1995). Other research suggests that δ15N depletion in trees may be related to fractionation during transfer from mycorrhizas and depletion of δ15N in tree shoots may indeed be indicative of mycorrhizal status (Hobbie et al., 1999, 2000). Problems of this kind must be borne in mind if the δ15N method is to be used. As a general rule, other indicators of nitrogen fixation such as nitrogen accumulation and the ability of the tree to nodulate should be ascertained before this method is used. Application of the 15N natural abundance method for measuring nitrogen fixation by trees is reviewed in detail by Boddey et al. (2000).

13.6 Methods Based on Nitrogen Fixation Transport Products Nitrogen is taken up from soil principally in the form of nitrate, whereas fixed nitrogen is assimilated into amides or ureides depending on the host legume before transport to the shoots. The proportion of nitrogen derived from nitrogen fixation at any given time can be estimated based on the relative proportion of the nitrogen in the xylem sap present as ureides (Peoples et al., 1989). Samples of bleeding sap from cut shoots are taken for analysis of total nitrogen and ureide nitrogen. This method can also be applied by extraction of dried petioles and determination of the ureide content, which makes sampling easier (Peoples et al., 1989). Ureide production appears to be limited to legumes of the tribes Phaseoleae, Indigoferae and Desmodieae (Giller, 2001), so that the only agroforestry species confirmed to transport ureides are Cordariocalyx gyroides and Desmodium rensonii (Herridge et al., 1996). Flemingia macrophylla belongs to the tribe Phaseoleae and may therefore transport ureides (Giller,

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2001), but this has yet to be confirmed. Comparisons of field estimates of nitrogen fixation in Cordariocalyx with the 15N natural abundance method showed close correspondence with estimates made using the ureide method (Peoples et al., 1996). Reports of ureide production in species within the tribe Robineae (such as Sesbania and Gliricidia) and the Mimoseae are almost certainly due to artefacts produced by coloured compounds during the analysis procedure. Attempts have been made to estimate nitrogen fixation in amide-exporting legumes using xylem sap analysis, but with less success (Peoples et al., 1989).

13.7 Estimating Total Amounts of Nitrogen Fixation With all of the above methods, biomass estimates and tissue nitrogen analyses are required to calculate actual amounts of nitrogen fixation, using these estimates of %N from nitrogen fixation. Although it can be seen that estimation of nitrogen fixation is difficult, there is still remarkably little quantitative information given the potential importance of nitrogen fixation in the long-term sustainability of agroforestry systems.

Chapter 14 Mycorrhizas D.L. GODBOLD

AND

R. SHARROCK

School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, UK

14.1 Synopsis Most trees and crop plants form mycorrhizal associations but our understanding of their role, particularly in mixed species associations, is still developing. Whereas trees form associations with both ectomycorrhizal (Fig. 14.1) and arbuscular mycorrhizal (or endomycorrhizal) fungi (see Fig. 15.4 on p. 295), herbaceous crop plants only form arbuscular mycorrhizas. There is strong evidence to show that mycorrhizal colonization can be of great benefit to plants but that effects of colonization vary markedly depending on environmental conditions and species of both host plant and fungus. The benefits to host plants are assumed to be

Fig. 14.1. Pinus strobus root with ectomycorrhizal colonization. © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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because mycorrhizal colonization may improve nutrient uptake, water relations, resistance to soil pathogens, and resistance more generally to adverse soil conditions. Furthermore, since mycorrhizas may infect many different host species, the mycelial network they develop below ground may connect plants and facilitate transfers of carbon and nutrients among them. In this chapter, the evidence for potential effects of mycorrhizal colonization is discussed, leading to the identification of key research questions and then description of the methods required to address them.

Importance of mycorrhizas for plant nutrition The improved mineral nutrition of mycorrhizal plants is well documented (Marschner, 1995). In particular, a role in the uptake of phosphorus by ectomycorrhizas or arbuscular mycorrhizas, and nitrogen uptake by ectomycorrhizas and to a lesser extent arbuscular mycorrhizas, has been described (George et al., 1995). In Eucalyptus grandis and Eucalyptus maculata, inoculation with the ectomycorrhizal fungus Pisolithus sp. enabled the seedlings to utilize organic nitrogen sources, which could not be utilized by non-mycorrhizal seedlings (Turnbull et al., 1995). In addition to the elements phosphorus and nitrogen, mycorrhizas have been shown to facilitate plant acquisition of magnesium (Jentschke et al., 2000), copper and zinc (Guo et al., 1996) and manganese (Marschner, 1995). The effects of arbuscular mycorrhizas on mineral acquisition of plants have recently been reviewed by Clark and Zeto (2000). Both ectomycorrhizas and arbuscular mycorrhizas can greatly increase the volume of soil exploited due to the extent and high surface area of the extramatrical mycelium. It has been shown in a number of investigations that the external hyphae of mycorrhizas can absorb phosphorus from outside the root depletion zone and transport it to the host plant. Some of the most convincing demonstrations of this have used experimental designs with separate root and hyphal compartments. In these investigations roots were excluded from the hyphal compartment by a 30 µm net which allowed mycorrhizal hyphae to pass. With such designs, in white clover (Trifolium repens) colonized by arbuscular mycorrhizas, a depletion zone of more than 11 cm due to phosphorus uptake by the extramatrical hyphae could be shown (Li et al., 1991). In an investigation with Norway spruce (Picea abies) and the ectomycorrhizal fungus Paxillus involutus, using a separate root and hyphal compartment, translocation of phosphorus by the extramatrical mycelium over a distance of 5 cm was demonstrated (Brandes et al., 1998). In Pinus sylvestris mycorrhizal with Suillus bovinus, phosphorus was translocated over 30 cm, mainly in rhizomorphs (Finlay and Read, 1986a). Effective uptake of phosphorus by the hyphae of both ectomycorrhizas and arbuscular mycorrhizas may also

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be because polyphosphates are accumulated in vacuoles, where they act as both a storage form of phosphorus and function in energy storage. In addition to the increase in surface area provided by the extramatrical mycelium, ectomycorrhizas have also been shown to exude organic acids and mobilize mineral phosphorus from sparingly soluble sources. The involvement of organic acid exudation in uptake of phosphorus in non-mycorrhizal roots is well documented (Marschner, 1995). The ectomycorrhizal fungus Paxillus involutus exudes high amounts of malate and oxalate (Lapeyrie et al., 1987; Marschner, 1994). Cumming and Weinstein (1990) demonstrated that Pinus rigida ectomycorrhizal with Pisolithus tinctorius was able to extract phosphorus from insoluble aluminium phosphate. Recently, it has been suggested that ectomycorrhizas are a primary factor in the weathering of minerals (Jongmans et al., 1997; Landeweert et al., 2001). Thus there is a strong possibility that organic acids are involved in the uptake of phosphorus and also other nutrients such as potassium, calcium and magnesium (Landeweert et al., 2001) by ectomycorrhizas. Both ectomycorrhizas and arbuscular mycorrhizas have high levels of phosphatase activity. In wheat (Triticum aestivum), roots infected with arbuscular mycorrhiza had higher levels of acid phosphatase than nonmycorrhizal roots (Tarafdar and Marschner, 1994). In Norway spruce ectomycorrhizal with a number of fungi, similar levels of acid phosphatase were found in mycorrhizal and non-mycorrhizal roots (Eltrop, 1993). However, in Norway spruce mycorrhizal with Thelephora terrestris, a higher phosphomonoesterase activity was found in mycorrhizal roots and rhizomorphs than in non-mycorrhizal roots. These results indicate that mycorrhizas have a high potential to mobilize organic phosphorus, but that there are differences between species of mycorrhizas. However, even if the levels of acid phosphatase activity are similar between mycorrhizas and non-mycorrhizal roots, the large surface area of the extramatrical mycelium will greatly increase the potential to mobilize organic phosphorus. There are few investigations of the contribution of mycorrhizas to the total uptake of phosphorus by a plant. In Norway spruce ectomycorrhizal with Paxillus involutus, 52% of the total phosphorus uptake was shown to be via the extramatrical hyphae (Brandes et al., 1998). In arbuscular mycorrhizal Trifolium repens, the hyphae contributed 70–80% of the total phosphorus uptake (Li et al., 1991). In a similar system with Trifolium repens, phosphorus uptake in non-mycorrhizal plants was only 3% of the uptake of mycorrhizal plants (Schweiger et al., 1999). In non-mycorrhizal Leucaena leucocephala, soil solution phosphorus concentrations 27–38 times higher were required for maximal growth than in mycorrhizal seedlings (Habte and Manjunath, 1987). These data indicate that mycorrhizas have a high potential to contribute to phosphorus acquisition by plants. It is

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possible that experimental systems overestimate the mycorrhizal contribution to phosphorus acquisition but, since the mycorrhizal contribution measured experimentally is so high, it is likely that under field conditions mycorrhizas play a very significant role in phosphorus acquisition.

Importance of mycorrhizas for plant water relations Both ectomycorrhizas (Lamhamedi et al., 1992) and arbuscular mycorrhizas (Allen, 1991) can improve the water relations of plants. For example, in two species of Acacia, Cornet and Diem (1982) showed that mycorrhizal plants had a higher number of open stomata under drought conditions. However, the mechanisms involved are still controversial. A number of mechanisms of increased host drought tolerance have been suggested, such as increased root hydraulic conductivity (Koide, 1985), alteration of stomatal regulation due to hormone signals (Allen et al., 1982; Barea and Azcón-Aguilar, 1982), osmotic adjustment (Augé et al., 1986), hyphal water transport (Hardie, 1985; Faber et al., 1991; Ruiz Lozano and Azcón, 1995) and improved phosphorus nutrition (Fitter, 1988). Improved water status has often been associated with improved host phosphorus nutrition (Nelsen and Safir, 1982; Graham and Sylvertson, 1984; Fitter, 1988). Mycorrhization nearly always results in an improved phosphorus nutrition and/or larger size of host plants, which often makes it difficult to deduce the mechanisms involved in improved water relations. However, a number of authors have shown that host water status is improved independently of host phosphorus nutrition (Augé et al., 1986; Bethlenfalvay et al., 1988; Peña et al., 1988; Sanchez-Diaz et al., 1990). Davies et al. (1992) increased the phosphorus nutrition of non-mycorrhizal plants by addition of fertilizers to produce pepper (Capsicum annum) plants mycorrhizal with Glomus deserticola and non-mycorrhizal pepper plants with similar levels of phosphorus. After drought acclimation, mycorrhizal plants were the most drought tolerant. These authors suggested that the extraradical hyphae might facilitate water uptake and improve drought resistance. Faber et al. (1991) have suggested a role of the hyphae in the transport of water. These authors suggested that hyphae of arbuscular mycorrhizas might have access to smaller pores within the soil and facilitate water uptake under drought conditions. To estimate the contribution of the extraradical hyphae to water uptake, techniques have been used with separate hyphal and root compartments, divided by a nylon screen. Using such techniques, George et al. (1992) found no evidence for the involvement of hyphae of arbuscular mycorrhizas in water transport of Agropyron repens. Using a similar experimental design, Ruiz Lozano and Azcón (1995) were able to show

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that, in association with lettuce (Lactuca sativa), extraradical hyphae of both Glomus deserticola and Glomus fasciculatum were involved in water uptake. However, differences in water uptake were shown for the two fungi under different water regimes. Under poor water supply Glomus deserticola transported a larger amount of water than Glomus fasciculatum, suggesting species differences in hyphal water uptake and transport.

Effects of mycorrhizas on plant–pathogen interactions Apart from beneficial effects on the nutrition and water supply of host trees, mycorrhizas have been proven to increase the resistance of trees to infection by pathogens of the fine roots (Marx, 1972; Duchesne et al., 1989). Different species of mycorrhizal fungi vary in their efficiency at preventing root infections, although the mechanisms involved in this process are rather poorly understood. Possible modes of action include the production of antibiotics by the mycorrhizal fungi, stimulation of host defence mechanisms and the physical barrier presented by the Hartig net in ectomycorrhizal roots (Marx, 1972). Gunjal and Patil (1992) showed that arbuscular mycorrhizal fungi increased the resistance of casuarina (Casuarina equisetifolia) to Fusarium vesicubesum. Infections of wheat with common root rot (Bipolaris sorokiniana) and root disease caused by Gaeumannomyces graminis and Fusarium culmorum were reduced by arbuscular mycorrhizal fungi (Thompson and Wildermuth, 1989; Innocenti and Branzanti, 1998). In the studies by Innocenti and Branzanti (1998), the colonization of the roots with Glomus did not reduce the disease index, but increased host tolerance to pathogen attack. Both ectomycorrhizal and arbuscular mycorrhizal fungi have been shown to produce antibiotics (Marx, 1969; Trofast and Wickberg, 1977). In addition, ectomycorrhizal fungi produce phenolic substances (Strobel and Sinclair, 1991) and oxalic acid (Duchesne et al., 1989), which suppress root pathogens. Arbuscular mycorrhizal fungi have also been shown to reduce damage to plants by sedentary endoparasitic nematodes (Borowicz, 2001). An ectomycorrhizal fungus (Pisolithus sp.) increased the growth of nematode-sensitive Acacia holosericea seedlings under glasshouse conditions and reduced the multiplication of the root-knot nematode, Meloidogyne javanica, possibly through the production of polyphenolic compounds (Duponnois et al., 2000).

Transfer of carbon and nutrients between plants Ectomycorrhizas and arbuscular mycorrhizas form a large mycelia network in the soil. As trees of different species may be colonized by the same

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species of ectomycorrhiza, and arbuscular mycorrhizas mostly have a wide host range, different species of trees and other plants may be connected by a common below-ground hyphal network. It has often been suggested that this hyphal network of mycorrhizas forms a pathway for the movement of carbon and nutrients between plants (Finlay and Read, 1986b). In forests with ectomycorrhizal trees a movement of carbon between trees has been shown. In laboratory studies, transfer of carbon has also been demonstrated between plants connected by an arbuscular mycorrhizal mycelium (Francis and Read, 1984). Although the transfer of carbon between species or between mature trees and seedlings of a species is of great ecological interest, in agroforestry and agricultural systems the transfer of nutrients, especially nitrogen and phosphorus, is more important. This is particularly the case for the transfer of nitrogen from nitrogen-fixing trees or crops to receiver plants. This invariably requires the movement of nutrients between arbuscular mycorrhizal plants. There are conflicting reports about the role of mycorrhizas in the transfer of nitrogen between plants (Hamel et al., 1991; Frey and Schüepp, 1993). Frey and Schüepp (1993) showed that the mycorrhizal mycelium enhanced the transfer of nitrogen between clover (Trifolium alexandrinum) and maize (Zea mays). In contrast, Hamel et al. (1991) found that the transfer between soybean (Glycine max) and maize was small. The transfer increased if there was direct contact between the roots. Between soybean and maize a greater transfer occurred for nitrogen supplied from fertilizer than nitrogen from biological fixation (Marschner, 1995). This suggests that plants restrict the movement of symbiotically fixed nitrogen. There is little evidence to show that there is a transfer of nitrogen from the plant to the fungus (Ikram et al., 1994). The direct transfer of nitrogen between plants may, however, be a function of the sink–source relationship between potential donor and receiver plants (Frey and Schüepp, 1993). Ekblad and Huss-Danell (1995) showed that in a mixture of Alnus incana and Pinus sylvestris the proportion of fixed nitrogen in P. sylvestris derived from A. incana was highest (9%) when P. sylvestris was nitrogen starved. Ikram et al. (1994) investigated nitrogen movement between Pueraria phaseoloides as a nitrogen-fixing donor plant and rubber trees (Hevea brasiliensis) as receiver plants. P. phaseoloides is used as a cover crop in rubber plantations. In the investigation, the shoot of the donor plant was shaded or removed to increase the difference in sink strength in favour of the receiver plant. Despite this treatment P. phaseoloides only supplied 0.07–0.4% of the total nitrogen uptake of H. brasiliensis. This suggests that mycorrhizal links play only a small role in the transfer of nitrogen in this system. However, it is known that intercropping of rubber with nitrogen-fixing P. phaseoloides, as in other intercropping systems, increases the growth of rubber (Broughton, 1977). The roots and mycorrhizas may take up nitrogen

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released by the decomposition of leaf and root litter. It has also been suggested that nitrogen may be transferred via root exudation of nitrogen compounds (see Table 13.1 on p. 262). Transfer from the roots may also be higher if the roots of the donor plants are dying. Johansen and Jensen (1996) found a significant transfer of nitrogen from pea (Pisum sativum) to barley (Hordeum vulgare) in mycorrhizal systems when, after removal of the shoot, the roots of the donor plant were dying. Mycorrhizas are able to take up and transport both inorganic and organic forms of nitrogen (George et al., 1995), both of which may be available in dying roots. Such a transfer may be of particular significance after pruning of nitrogen-fixing trees in agroforestry systems, and in trees with high rates of root turnover.

Differences between mycorrhizal species and strains In an investigation of the distribution of arbuscular and ectomycorrhizal tree species in savannas, Högberg (1989) suggested that ectomycorrhizal trees dominate soils with low levels of nitrogen and phosphorus, whereas arbuscular mycorrhizal species dominate soil low in phosphorus but relatively rich in nitrogen. This suggests that ectomycorrhizal trees may have a greater capacity for nitrogen acquisition than arbuscular mycorrhizal species. However, in the humid tropics of French Guiana, ectomycorrhizal tree species were found not to dominate the poorest soils (Bereau et al., 1997). The considerable benefits of mycorrhization can be seen in the advantages conferred by efficient species and strains. There are, however, significant differences between the effects that different species and strains of both ectomycorrhizas and arbuscular mycorrhizas have on host plants. It has been shown that the potential to take up phosphorus varies strongly between species both for ectomycorrhizas and arbuscular mycorrhizas (Marschner, 1995). As described above, species differences have also been demonstrated for improved water relations and pathogen resistance. Similarly, there are considerable functional differences between species of mycorrhizas and different species of trees. Bâ et al. (2000) demonstrated that only five species of tropical fruit trees (Zizyphus mauritiana, Tamarindus indica, Dialium guineensis, Parkia biglobosa and Cordyla pinnata) out of a total of 15 species had higher dry weight as a result of inoculation with Glomus aggregatum. In Zizyphus mauritiana and Cordyla pinnata a smaller positive effect was shown after inoculation with Glomus intraradices. At present there is a potential danger of either underestimating or overestimating the significance of mycorrhizas because only a small number of mycorrhizal species have been investigated. For example, to reach reliable estimates of the contribution of mycorrhizas to phosphorus

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uptake by plants in the field, we need to know the species composition of the mycorrhizas colonizing roots and have information about the phosphorus uptake capabilities of each species. The development of molecular biological methods of species identification for both ectomycorrhizas and arbuscular mycorrhizas has greatly increased our understanding of species composition (see Section 14.5).

Inoculation and management of mycorrhiza populations Considering the advantages conferred by mycorrhizal infection, it is easy to understand why inoculation with mycorrhizas normally greatly increases growth and survival of host plants over that of non-mycorrhizal plants. Increased growth as a result of inoculation with mycorrhiza has been shown on numerous occasions in plants used in agroforestry systems (Osonubi et al., 1991). However, the majority of studies that have investigated the advantages of mycorrhizal infection have compared mycorrhizal plants with non-mycorrhizal plants, which is unrealisitic since in most soils, with the exception of soils with an extremely low inoculation potential, most plants will form some sort of mycorrhizal association albeit weak. However, with an increasing knowledge of the importance of mycorrhizal species differences, there is an increasing interest in the introduction of mycorrhizal species that are more efficient than the species present in a given soil under field conditions. Once introduced, these mycorrhizas must persist in the soil together with the indigenous mycorrhizas. Due to the difficulties in species identification of mycorrhizas, especially arbuscular mycorrhizas, there are few studies to estimate the persistence of introduced mycorrhizal species. With the development of better identification methods a clearer picture of mycorrhizal persistence will emerge. It is reasonable to assume that the highest success rate for introduced mycorrhizas will be achieved in soils with the least competitive indigenous mycorrhizal flora. However, even if introduced mycorrhizal species have a low persistence, the advantages conferred on the plant during establishment may be sufficient to justify their use. This may be particularly important for trees under adverse conditions, where establishment is often the biggest problem. Management options for mycorrhizal populations in agricultural systems have been reviewed several times (Barea and Jeffries, 1995; Haselwandter and Bowen, 1996). Briefly these include using crop rotation techniques, precropping or intercropping. In each case a plant species is used to maintain a high inoculum potential in the soil. For example, a high level of mycorrhizal infection was maintained in cassava (Manihot esculenta) when it was grown in rotation with groundnut (Arachis hypogaea) (Sieverding and Leihner, 1984). Dodd et al. (1990a,b) showed that

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precropping with cassava, kudzu (Pueraria phaseoloides) or sorghum (Sorghum bicolor) significantly increased arbuscular mycorrhizal infection of cowpea (Vigna unguiculata). As intercrops, millet (Pennisetum glaucum), Sudan grass (Sorghum sudanense) and sorghum increased the population densities of arbuscular mycorrhizal fungi and improved the growth of tree seedlings (Johnson and Pfleger, 1992). The reverse may also be true where, in agroforestry systems, highly mycorrhizal trees may be used to maintain the inoculum potential in the soil. Roots of Citrus aurantifolia grown in low-input systems had a higher degree of mycorrhizal colonization than those grown in conventional high-input systems in Mexico (Michel-Rosales and Valdés, 1996). These effects may have been related to fertilizer input, but the low-input systems were also structurally diverse, with a large number of other species including 16 species of fruit trees, compared with the high-input systems, which were monocultures. However, on a Nigerian Alfisol, Atayese et al. (1993) found that Senna sp., Gliricidia sp. and Leucaena sp. did not increase the mycorrhizal spore density of the soil even though Gliricidia sp. and Leucaena sp. were highly mycorrhizal. The mycorrhizal spore density was, however, increased by planting cassava (Manihot esculenta).

Research needs Many of the studies cited above have been carried out under laboratory conditions. Therefore, there is a general requirement to transfer results obtained in laboratory experiments to the field. Many of the questions that need to be answered for mycorrhizas in tree–crop associations, listed below, are similar to the questions relevant for conventional agriculture and forestry. •

•

Considerably more information is required about the ecophysiology and effectiveness of different mycorrhizal species, as well as their below-ground population dynamics. Information about species should include estimates of which nutrient pools mycorrhizas are accessing and whether these differ from non-mycorrhizal plants. Information is also required about the persistence of introduced mycorrhizas, and whether persistence can be increased by cultivation techniques. Additionally, information is required on how management actions, such as mulching, crop rotation and fertilization (both mineral and organic), affect the development of mycorrhizas.

In addition there are some research questions specific to tree–crop associations.

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Can the association of crops with strongly mycorrhizal trees or tree species with particularly efficient mycorrhizas increase the access of these crops to less available nutrient pools in the soil via nutrient release from tree litter? Can strongly mycorrhizal species aid the establishment of poorly mycorrhizal species under adverse site conditions, for example in rehabilition of degraded lands?

14.2 Inoculation Methods Propagules of mycorrhizal fungi include soil hyphae, spores, old root fragments and colonized particles of organic matter (Brundrett and Abbott, 1994). In particular, soil hyphae are important propagules under drought conditions. Soil disturbance and degradation often result in low inoculation potential of the soil; hence there is often a need to reintroduce mycorrhizal fungi in general or to introduce efficient mycorrhizal strains or species. Land management practices can then be used to maintain introduced mycorrhizas or enhance indigenous populations. The simplest source of mycorrhizal propagules is soil from areas of high mycorrhizal activity. Soil may be used as a non-specific inoculum for both ectomycorrhizal and arbuscular mycorrhizal fungi. However, for reasons of phytosanitation it is often undesirable to use soil as inoculum. For this reason and also to allow introduction of specific fungi, artificial inocula are often required. For ectomycorrhizas, inoculation techniques have been developed which use spores of selected fungi (Marx et al., 1984). Mycelium of ectomycorrhizal fungi can easily be produced in fermenters. The mycelium can then be trapped in alginate beads for use as inoculum (Mauperin et al., 1987). Arbuscular mycorrhizal fungi cannot be cultured ex planta and thus must be cultured with a host plant. Most inoculum production requires growth of the host plants in a medium that can be used as a carrier for spores and hyphae. Carrier substances include expanded clays (Feldmann and Idczak, 1994) or vermiculite or perlite (Bagyaraj, 1994). Soil-free inoculum has been produced in aeroponic (Jarstfer and Sylvia, 1995) and nutrient-film cultures (Elmes and Mosse, 1984). The suitability of these inocula for different applications depends on the identity of the main mycorrhizal propagules and their ability to retain infectivity during storage and to persist in the soil from year to year, as well as on methods available for application (Smith and Read, 1997). Due to the expense of the production of inoculum and the bulk required, large-scale inoculation is often not a viable option. Thus management of indigenous populations is of great importance (see Section 14.1).

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14.3 Sampling of Plant Roots for Mycorrhizal Studies Collection of plant roots for estimating mycorrhizas requires sampling, preparation and storage of the roots. Root samples are normally collected using a trowel or soil auger (see Section 12.2). Roots are normally removed from the soil using careful washing with water or removal of the soil using forceps. Particularly for visual identification of ectomycorrhizas damage to the roots should be avoided. For quantification and identification (both visual and molecular) of arbuscular mycorrhizas and for molecular identification of ectomycorrhizas, roots can be stored in 50% (v/v) ethanol. For visual quantification and identification of ectomycorrhizas, roots can be stored in moistened Petri dishes at 4°C for several days. Fixation of ectomycorrhizas often results in colour changes, making visual identification more difficult.

14.4 Quantification of Mycorrhizas Non-vital staining of arbuscular mycorrhizas In order to visualize and quantify root colonization by arbuscular mycorrhizal fungi the fungal structures must be stained. The most commonly utilized stains are non-vital stains, which cannot distinguish between living and dead fungal tissues. Stains for light microscopy include trypan blue (Koske and Gemma, 1989) and chlorazol black E (Brundrett et al., 1984). Acid fuchsin is also a popular non-vital stain; this stain has fluorescent properties, which are best visualized by fluorescence microscopy (Merryweather and Fitter, 1991). One of the most frequently cited non-vital staining procedures utilizes the fungal staining properties of trypan blue (Phillips and Hayman, 1970; Koske and Gemma, 1989). Roots are washed from the soil and either stained directly or fixed in 50% ethanol and stained at a later time. The root tissue is cleared in 2.5% KOH at 90 °C for 10–30 min in a water bath. After clearing, dark tissues may need to be bleached in a freshly prepared solution of alkaline hydrogen peroxide for 10–45 min. Following the KOH and bleaching steps, the root tissue is highly alkaline and requires acidification prior to staining with trypan blue. The roots are rinsed thoroughly and acidified in 1% HCl for 1–24 h (at least 8 h is generally recommended). The acidified roots can then be stained in 0.05% trypan blue in acidic glycerol at 90 °C for 15–60 min. The stained tissues are then de-stained in acidic glycerol at room temperature. De-staining overnight generally improves the contrast between the fungal and root tissues, which often initially retain excess stain. Stained tissues can be stored in acidic glycerol in the dark for

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many months. For the protocol details in full see Koske and Gemma (1989).

Visual methods of evaluation of arbuscular mycorrhizas The assessment of fungal colonization is generally based on estimates of the proportion of potential host tissue (the root cortex) colonized by arbuscular mycorrhizal fungi (Giovannetti and Mosse, 1980; McGonigle et al., 1990). The most extensively used visual method of determining fungal colonization is the grid-line intersect method (Giovannetti and Mosse, 1980; McGonigle et al., 1990). This method is used to estimate the proportion of colonized tissue and total root length. A root sample is spread across a grid of known dimensions drawn on the base of a Petri dish. Using a dissecting microscope at ×40 magnification, each point at which a root intersects a vertical or horizontal grid line is studied and the presence or absence of fungal tissue is noted. However, at ×40 magnification it is difficult to distinguish between different fungal structures. In light of this McGonigle et al. (1990) devised an objective method of measuring root colonization called the magnified intersection method. This method is used to obtain the percentage colonization of different fungal structures in a root system. Root sections, approximately 1 cm in length, are mounted on microscope slides in a few drops of acidic glycerol and viewed with a compound microscope at magnification ×200. Root/eye-piece cross-hair intersections are considered along the axis of the root section and the presence of arbuscules, hyphae and vesicles is noted at each intersection. A fixed number of intersections are considered per root section. From these data the percentage arbuscular, hyphal and vesicular colonization can be calculated.

Visual methods of evaluation of ectomycorhizas The degree of colonization of ectomycorrhizas can be estimated by counting the number of mycorrhizal and non-mycorrhizal root tips in a representative sample (>100 root tips).

Vital staining To determine the proportion of active fungal colonization associated with a root, vital staining procedures are employed. Vital stains locate sites of enzymatic activity, which are specific to viable fungal structures. Nitroblue

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tetrazolium is a vital stain which is coupled to the activity of succinate dehydrogenase (a tricarboxylic acid enzyme), to produce formazan, which is purple (Smith and Dickson, 1991; Schaffer and Peterson, 1993). Used in conjunction with a non-vital stain such as acid fuchsin, the proportion of active to total colonization can be assessed. Alkaline phosphatase activity is also considered to be a good indicator of fungal viability. Alkaline phosphatase is sequestered in the phosphataseaccumulating vacuoles in hyphae and is also found along the fungal tonoplast, but is not present in root tissue (Dickson and Smith, 1998). Other vital (and non-vital) stains are reviewed by Dickson and Smith (1998).

Laser scanning confocal microscopy Visualization of mycorrhizal structures and quantification of their surface area and volume have recently been achieved by laser scanning confocal microscopy after staining with acid fuchsin (Dickson and Kolesik, 1999).

Chitin assay The amount of fungal tissue present in the host tissue can be measured by chemical means. The chitin assay is based on the quantitative determination of chitin, which is present in the fungus, but not in the host. Total chitin is measured by colorimetric assay after conversion into glucosamine (Hepper, 1977; Bethlenfalvay et al., 1981).

Molecular quantification Quantification of endomycorrhizal fungi in a root system has also been achieved by molecular means using an assay based on competitive polymerase chain reaction (PCR) (Edwards et al., 1997).

14.5 Identification of Mycorrhizal Fungi Visual identification Many ectomycorrhizas can be identified using a visual method developed and described by Agerer (1992, 1993). This method involves using both morphological features and the hyphal mantel characteristics to identify to genus or species level. Visual methods can also be used to identify some

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arbuscular mycorrhizas; however, this method is of limited potential as a single mycorrhizal species may have a different morphology between host plant species.

Immunological approaches Immunochemical detection methods are based on the reaction between antibodies and antigens. The production of antibodies is an immune system response to alien substances (the antigen). The type of antibody produced is often antigen-specific. It is this specific relationship between antibody and antigen that can be utilized as a tool in fungal identification (Göbel et al., 1998; Hahn et al., 1998). Antibodies, produced by exposing an animal’s immune system to a specific antigen, are coupled by chemical means to molecules which can be easily detected, such as fluorescent dyes for fluorescence microscopy, enzymes for enzyme-linked immunoassays, or heavy metals for immunocytochemical analysis with an electron microscope (Harlow and Lane, 1988; Göbel et al., 1998; Hahn et al., 1998). Labelled antibodies introduced into a system in which the fungal symbiont is unknown will recognize and bind to their corresponding antigen (providing it is present). Schmidt et al. (1974) were among the first to analyse mycorrhizal fungi by immunochemical means. More recently, polyclonal antibodies have been generated with sufficient stringency to detect Scutellospora species in the roots of several plants (Thingstrup et al., 1995). Dot immunoblots have also been used in an attempt to identify Gigaspora and Acaulospora species (Sanders et al., 1992).

Isozyme analysis Isozymes are different molecular forms of the same enzyme. They generally have the same enzymatic properties and differ only in their amino acid composition and net charge. Due to these differences in net charge, isozymes can be differentiated by their electrophoretic mobility (Rosendahl and Sen, 1994). Many isozymes are apparently species-specific in nature. Tisserant et al. (1998) ran protein extracts taken from root tissue colonized by five Glomus species on a polyacrylamide gel. The separated protein bands were stained for ten different enzymes, including alkaline phosphatase and malate dehydrogenase. Different banding patterns emerged, revealing the presence of several mycorrhizal-specific isozymes. These mycorrhizal-specific isozymes were subsequently successfully used as fungal-specific markers in plants grown in field trials. Isozyme analysis is also increasingly used in conjunction with PCRand restriction fragment length polymorphism (RFLP)-based molecular

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approaches to identifying both endomycorrhizal (Dodd et al., 1996) and ectomycorrhizal (Timonen et al., 1997) fungi in root tissue.

Mycorrhizal identification by molecular means Molecular approaches to mycorrhizal identification are becoming increasingly popular. Wyss and Bonfante (1993) extracted genomic DNA from the spores of Glomus versiforme and Gigaspora margarita isolates. Genomic fingerprints of these endomycorrhizal isolates were generated by PCR amplification with short arbitrary primers, a process known as random amplified polymorphic DNA (RAPD-PCR). The RAPD-PCR method employs an arbitrary 10 base-pair primer, which anneals to a number of complementary sites on the template DNA. The PCR products generated are separated according to their molecular weights on an agarose gel and stained with ethidium bromide. The banding pattern on the gel reflects the overall structure of the DNA molecule used as the template. If the starting material is total cell DNA, then the banding pattern represents the organization of the genome (Williams et al., 1990). Wyss and Bonfante (1993) found that different mycorrhizal species generated different banding patterns. To a lesser extent, differences were also observed between different isolates of the same species. Isolate-specific RAPD-PCR bands have been used to generate isolatespecific PCR primers by Abbas et al. (1996). The DNA present in bands unique to isolates of Glomus mosseae and Gigaspora margarita was purified, cloned and sequenced. Sequence analysis subsequently led to the generation of isolate-specific primer pairs, which in conjunction with PCR technology was utilized to identify these isolates in root systems. The presence of a particular isolate in a root system is identified by the presence of a PCR product band on an agarose gel. Ribosomal DNA (rDNA) sequences have also been extensively utilized in the generation of mycorrhizal-specific PCR primers. Using PCR, RFLP, cloning and sequencing technologies, sequence differences have been highlighted in the following rDNA regions: the small subunit (SSU)/18S gene, the large subunit (LSU)/28S gene, internal transcribed spacer (ITS) regions and intergenic spacer (IGS) regions (see Egger, 1995). These differences are apparently family/species/isolate specific and have been used to generate mycorrhiza-specific PCR primers. Simon et al. (1993) used single subunit (SSU)/18S gene sequences from a number of endomycorrhizal fungi to generate family-specific primers. The SSU sequences were aligned and regions apparently unique to a family were selected for the design of family-specific primers. Members of the Acaulosporaceae, Gigasporaceae and Glomaceae have been successfully identified using these family-specific primers in spores and root systems.

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Van Tuinen et al. (1998) utilized the LSU/28S rDNA region to generate species/isolate-specific primers for the endomycorrhizal fungi Glomus mosseae, Glomus intraradices, Scutellospora castanea and Gigaspora rosea. Comparable studies have been carried out on endomycorrhizas by Sanders et al. (1995), Clapp et al. (1999) and Helgason et al. (1999). Ectomycorrhizas have also been subject to molecular analysis. Erland (1995) studied the abundance of Tylospora fibrillosa ectomycorrhizas in a spruce forest in Sweden. Tylospora fibrillosa could be distinguished from a large number of other basidiomycetes by the banding pattern generated by PCR amplification of the rDNA ITS region followed by RFLP analysis of the PCR product. Paolocci et al. (1999) also utilized the rDNA ITS region to identify Tuber species, which were successfully characterized on host plants by PCR amplification using species-specific ITS primers. The analysis of rDNA has also been used to compare the ericoid mycorrhizas Scytalidium vaccinii and Hymenoscyphus ericae (Egger and Sigler, 1993).

Indirect methods of quantification using spores and sporocarps The population structure of arbuscular and ectomycorrhizas can be estimated using soil spores for arbuscular mycorrhizas (Tommerup, 1994), or sporocarps (mushrooms) for ectomycorrhizas. The spores of arbuscular mycorrhizas must be isolated from the soil (Pacioni, 1994) and can be identified to genus or species level using morphological characteristics (Brundrett et al., 1996). For ectomycorrhizas, sporocarps within an area can be collected, identified and their abundance estimated. However, the spore and sporocarp production is dependent upon a number of seasonal factors (temperature, moisture), and often correlates poorly with the below-ground abundance (Tommerup, 1994; Erland, 1995). Additionally, a number of important ectomycorrhizas do not form sporocarps.

14.6 Estimation of Mineral Nutrient Uptake Through Mycorrhizas Absorption, translocation and transfer of mineral nutrients by the external hyphae of mycorrhizas have been demonstrated a number of times using radioisotopes (Tinker et al., 1994). Most of the attempts at quantification of mineral nutrient uptake by mycorrhizal fungi have used techniques in which separate root and hyphal compartments have been established using fine mesh (Li et al., 1991; George et al., 1995; Schweiger et al., 1999; Jentschke et al., 2000). The size of the mesh is usually between 20 and 45

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µm, which allows mycorrhizal hyphae to penetrate, but restricts growth of roots. The proportion of mineral nutrients taken up by the mycorrhizas is normally estimated by determining the decrease in mineral elements in the soil (Li et al., 1991), or by using radioisotopes (Schweiger et al., 1999) or stable isotopes (Jentschke et al., 2000) as tracers.

Transfer of nitrogen from nitrogen-fixing plants Techniques to measure the transfer of nitrogen from nitrogen-fixing plants to non-nitrogen-fixing plants via mycorrhizal hyphae have used compartments to prevent root contact as described above (Ikram et al., 1994; Ekblad and Huss-Danell, 1995). A mesh is used to allow only mycorrhizal hyphae of the donor plant to be in contact with roots or hyphae of the receiver plants (Fig. 14.2). In some cases legumes grown in split root cultures have been used (Reeves, 1992). Nitrogen transfer is estimated using 15N as a tracer. 15N-labelled fertilizers are applied to the donor plants, and the amounts of fixed and transferred nitrogen is calculated using 15N-dilution methods (Giller et al., 1991). 15N is determined by mass spectrometry. 20–45 µm mesh

Mycorrhizal hyphae

Fig. 14.2. Experimental set-up used to investigate transfer of nitrogen from nitrogen-fixing plants to non-nitrogen-fixing plants via mycorrhizal hyphae. Root and hyphal soil compartments are separated by a fine mesh of pore size 20–45 µm, which allows passage of hyphae but not roots.

Chapter 15 Rhizosphere Processes D. JONES School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK

15.1 Synopsis The rhizosphere can be defined as a zone of intense biological and chemical activity in the soil that surrounds the root. The rhizosphere is chemically, physically and biologically different from the bulk soil because it has been modified by roots and their associated microorganisms. Many key processes associated with soil fertility occur in this modified soil and the rhizopheres of different plants may interact. Where there are very high root densities, as in silvopastoral systems, most of the top 15–20 cm of soil may be modified by roots in this way, so that it all effectively constitutes rhizosphere soil. A comprehension of the rhizosphere is fundamental to understanding the interactions of plants with their environment and their ability to respond to stress. Different plants modify the environment around their roots in different ways and, therefore, interactions between plants may occur as a result of rhizosphere processes. The key rhizosphere processes that affect interactions between trees and soil fertility can be classified under three headings: • •

•

chemical processes, especially those affecting nutrient capture (see Chapter 8) and metal toxicity (see Chapter 5); biological processes, including symbiotic associations (see Chapters 13 and 14), pathogenic activity and allelopathy (where the release of chemicals from roots of one plant affect other plants); and physical processes, including modification of soil structure (see Chapter 10) and its impact upon water availability (see Chapter 11).

© CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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The vast majority of research that is done on plant interactions with soil in agroforestry focuses on gross effects of, and changes in, soil properties, rather than the detailed rhizosphere mechanisms through which these come about, but it is necessary to understand basic processes if results are to be generalized and extrapolated. Much rhizosphere research has concentrated on only a few model plant species, in very controlled conditions, and its relevance to tropical agroforestry is unclear. This means that there is considerable scope to extend rhizosphere research to determine underlying mechanisms governing interactions in economically important tree and crop associations that are used in agroforestry.

Potential importance of rhizosphere processes in agroforestry Agroforestry systems employ two or more, often contrasting, plant species such as shade trees and herbaceous grasses in silvopastoral systems, or cereal crops and nitrogen-fixing trees in hedgerow intercropping. The premise of these systems is that the interactions between associated plants with respect to below-ground resources such as nutrients and water are complementary or facilitative rather than competitive (see Section 5.1). In some cases, one of the species may be partly chosen to stimulate aboveand below-ground nutrient cycling (typically of phosphorus and nitrogen), as in the case of leguminous cover crops, legume shade trees and planted fallows. The quantification of this enhanced cycling can be achieved at a gross scale using stable isotopes and radiolabelled tracers (13C, 15N, 32P, 33P, 35S; see Sections 7.3 and 8.2). However, this provides little mechanistic information. In some cases it can be shown that co-planted species have different rooting strategies that result in nutrients being taken up from different spatial zones in the soil, but in other cases rooting patterns do not provide a mechanistic basis for the complementary or competitive effects observed and the mechanisms involved can only be speculated upon (Kamara et al., 1999; Lehmann et al., 1999a; Ndufa et al., 1999). To fully understand why these effects occur requires an understanding of nutrient flows in the rhizosphere. For example, where enhanced phosphorus cycling is observed, the impact of both mycorrhizal and root phosphorus mobilization strategies needs to be compared in each species to identify the spatial and temporal mobilization of different soil phosphorus pools such as organic phosphorus, iron–aluminium hydroxide-fixed phosphorus and calcium-bound phosphorus (see Section 5.3). Furthermore, the plant species chosen for agroforestry systems may form different mycorrhizal associations (see Chapter 14). Trees may be either endo- or ectomycorrhizal or both, such as many species of the Eucalyptus genus, whereas associated crops may only be endomycorrhizal,

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as is the case with maize, or may not form mycorrhizal associations at all, such as Brassica spp. It can be envisaged that under these circumstances the mycorrhizas will develop vastly different rhizospheres leading to contrasting patterns of nutrient capture, although experimental evidence for this form of complementary resource use is lacking. In situations where the woody plant has the same mycorrhizal association as the crop, such as a Tithonia diversifolia hedge next to a maize crop, then competition for soil resources may be more intense. However, as the rhizosphere of each plant species can be assumed to be unique, it is likely that resource capture by plants will always differ to some extent. The presence of one root system, such as that of a tree, can also aid in the resource capture by a crop plant. This is evidenced by both the direct and indirect transfer of nitrogen from nitrogen-fixing plants to nonnitrogen-fixing plants (see Chapter 13). In addition, a number of other interactions may occur where trees with abundant mycorrhizas can release root exudates capable of mobilizing the large reserves of organic nitrogen or phosphorus, making it available for associated crops. While our knowledge of the rhizosphere characteristics of different plant species remains so sparse we can only speculate upon the many interactions that may occur in species associations. Physical attributes of soil are also modified by roots in the rhizosphere and these strongly influence the growth and activity of organisms living in soil. Root exudates have been shown to significantly improve soil structure (see Chapter 10). This can occur directly in response to the root addition of polysaccharide gel (mucilage) to the rhizosphere, which helps bind aggregates together and also enhances soil–root contact (Czarnes et al., 2000). Indirectly, the release of both high and low molecular weight root exudates can also enhance soil structural stability through the stimulation of the rhizosphere microbial community, which in turn produces polysaccharide gels (Traore et al., 2000). As plants release different amounts of root exudates it can be expected that they influence soil physical properties in different ways. This is of relevance in an agroforestry context, where the maintenance or rehabilitation of soil structure is required and so plants may be selected specifically for their ability to enhance soil structure. Exudation of moisture by roots into dry rhizosphere soil can also be significant in wetting soil and so facilitating nutrient uptake by the plant whose roots exude the water as well as nutrient and water uptake by other plants in the vicinity (see discussion of hydraulic lift in Section 11.1). This clearly has implications for nutrient capture from otherwise dry soil and both competitive and facilitative interactions among species with different rooting habits.

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Conceptual and methodological difficulties of rhizosphere studies Until recently, it has been difficult to achieve a mechanistic understanding of below-ground interactions between plants and the importance of rhizosphere processes in these interactions because techniques have not been available for unravelling the complexities of the processes involved. Indeed, we are still far from gaining a holistic understanding of the rhizosphere as we are only able to gain insights into specific areas for which techniques are available. Although there is conclusive evidence that the rhizosphere exists, its spatial boundaries remain difficult to define. This means that when embarking on rhizosphere research, a key step is to define an effective rhizosphere for the purposes at hand. In the case of chemicals lost from the root it is known that some may have an extensive diffusion sphere of many centimetres, whereas other highly charged compounds may only travel a few micrometres away from the root (Darrah, 1991a). Although they differ according to the chemical involved, the vast majority of chemical processes in the rhizosphere occur within a few millimetres of the root (Fig. 15.1). The exact definition of the rhizosphere and consequently the volume of soil that needs to be studied has to be relevant to the hypotheses being tested. For example, when studying allelopathic interactions in the rhizosphere, the effective radius of the rhizosphere is controlled by the diffusion of the allelopathic chemicals.

Mineral nutrients

Organic carbon

1.00

600

NO3–0.50

0.25

Insoluble C –3

K

0.75

C concentration (µg cm )

Concentration after 10 days Concentration in bulk soil

P

450

Soluble C 300

150

Microbial biomass C

0.00 1.00

1.25

1.50

1.75

Distance from root surface (mm)

2.00

0 1.00

1.25

1.50

1.75

2.00

Distance from root surface (mm)

Fig. 15.1. Mineral nutrient and carbon profiles in a rhizosphere after 10 days according to simulation models based on experimentally derived nutrient uptake rates of maize and wheat. The mineral nutrient concentrations are relative to the amount present in the bulk soil at day 0 (adapted from Barber, 1995, and Darrah, 1991b).


Root cap mucilage

Epidermal cell mucilage

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Bacterial cell mucilage Epidermal cell death and loss

Root cap

Sloughed off root cap cells

Cortical cell death and loss

Wound site (e.g. soil particle abrasion damage, where a secondary root emerges)

Fig. 15.2. Different functional zones of a root; root hairs are not shown.

Leaving the boundaries aside, there is also a high degree of both spatial and temporal complexity. Questions therefore arise as to whether roots should be treated as whole units or subdivided based on size, age or functional zone (Fig. 15.2). For example, root hairs (not shown in Fig. 15.2) can be usefully distinguished from the root apex. It is certainly clear from a root physiology point of view that the behaviour of different root areas is very different (Marschner, 1995). These functional root units can arise because of localized suberization/lignification, the degree of apoptosis or other forms of cell death, the different water and chemical status of roots in different soil environments or the degree of mycorrhizal infection. It is for this reason that most rhizosphere experiments are conducted in laboratory mesocosms under optimal growth conditions rather than in the field, where spatial heterogeneity makes it difficult to interpret results. Typically, laboratory rhizosphere experimentation involves the use of seedlings and crop plants such as wheat, maize and bean. To some extent this has set a precedent so that the body of knowledge is so great for these model plants that further research tends to pursue research on these to the detriment of other species. The role of rhizosphere processes in the ecology of non-crop plants is almost unknown apart from a few isolated examples (Jones, 1998). In addition, work on trees has also been extremely limited, mainly focusing on temperate trees typically no more than a few years old due to difficulties in managing root mycorrhizal status, space limitations in climate control chambers and the short time horizon of much research (Grayston and Campbell, 1996; Grayston et al., 1997). Despite all the limitations on performing rhizosphere experiments, significant technical advances have been made in recent years which have allowed us to gain new insights into the biological interactions below ground. Although most of these techniques rely heavily on laboratory experimentation using either mesocosms (volume c. 50–2000 cm3) or more

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export

Soil atmosphere

Root

Shoot import

fixation

mineralization

efflux

mass flow and diffusion

uptake sorption and fixation

efflux

Microbial biomass

Soil mineral particles

Soil solution desorption and dissolution

uptake

death

mineralization

Necromass

incorporation

sorption and fixation

breakdown and dissolution

Soil organic pool

Fig. 15.3. Schematic representation of a mechanistic flux framework describing some of the possible nutrient fluxes in the rhizosphere.

often microcosms (volume c. 5–50 cm3), these should not limit their potential use in agroforestry research. Although this necessitates a more reductionist approach it also reinforces the need for large multidisciplinary projects and, in the case of trees, long-term research funding. Many of the methods outlined below were first developed for looking at biological interactions in bulk soil but have since been adapted for use in small soil volumes such as the rhizosphere. One of the major limitations to many rhizosphere studies performed in the past has been a lack of understanding of the complexity of the rhizosphere and an ignorance of key variables. For example it is common to investigate exudate release from roots without considering microbial consumption of the exudates (Matsumoto et al., 1979). Where experiments are aimed at parameterizing models, it is necessary in many cases to develop a mechanistic flux framework. An illustration of this is given in Fig. 15.3. This conceptualization provides a basic decision-making tool for ensuring that all factors are being considered in order for the results to be interpretable.

15.2 Obtaining a Representative Sample of Rhizosphere Soil Rhizosphere soil can be arbitrarily defined as the sheath of soil remaining attached to the roots after removal from the ground and gentle shaking. This offers a practical and much used approach to an almost impossible task. However, the amount of soil recovered can be highly dependent on soil and plant conditions. Soil is held to the root via a number of


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Table 15.1. Comparison between root sap and soil solution nutrient concentrations illustrating the need not to contaminate rhizosphere soil samples with plant sap. Solute concentration (mM)

K+ Ca2+ NO–3 PO34– SO42– Simple sugars Amino acids Organic acids

Root sapa

Soil solutionb

Fold difference

100 8 30 7 6 50 10 10

0.1 4 1.2 0.001 0.2 0.05 0.1 0.05

1000 2 25 7000 30 1000 100 200

aZea

mays root sap (includes cytoplasm and vacuolar sap). solution extracted by centrifugal drainage from a free-draining Typic Haplorthod.

bSoil

mechanisms including binding by water, fungal hyphae (mycorrhizal and non-mycorrhizal), adhesion with root and bacterial polysaccharides (mucilages) and adhesion via root hairs. Of these factors, soil moisture appears to largely determine the amount of soil recovered by this method, with little soil recovered when the soil becomes either too wet or too dry. There is, therefore, a critical moisture balance required for obtaining intact soil rhizosheaths and, where possible, soil moisture should be normalized across treatments. The effectiveness of the method is further reduced because soil adhesion is not usually uniform along the root length, making

Bulk soil

SOIL

Mycorrhizal hypha

Ectorhizosphere

Bacterial colony Root and bacterial mucilage

Rhizoplane

Root epidermal cell

Mycorrhizal arbuscule

ROOT

Root cortical cell

Endorhizosphere

Fig. 15.4. Schematic representation of the rhizosphere zones associated with a vesicular–arbuscular mycorrhizal-infected root.

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comparisons between root zones difficult. It is unusual, for example, for soil to adhere to the root apex. Once this stage has been achieved, the subsequent separation of rhizosphere soil from the root is also difficult in that the two are inextricably linked and recovery of one is impossible without some degree of contamination by the other. This is particularly important in soil or root samples that are being used for nutrient (organic and inorganic) and microbial analysis (enzymes or chemical biomass measures) as root sap solute concentrations are typically much greater than those in the soil solution (Table 15.1). Further, the most intense site of rhizosphere activity, the rhizoplane, which is the root surface, and the endorhizosphere, which is the root apoplast, are rarely recovered by this technique (Fig. 15.4). To obtain non-contaminated and carefully controlled samples of rhizosphere soil typically requires a scaling down of experiments to a mesocosm or microcosm scale, that is, to pots or rhizotrons. Although this does not obviate performing experiments in the field, most studies performed to date have been carried out within climate-controlled chambers, enabling dismantling in the laboratory (Marschner, 1995). Briefly, roots are kept separated from rhizosphere soil through a series of nylon or polypropylene meshes. The choice of membrane is dependent on root and root hair size; however, two types of mesh are usually employed. The first is a 0.2 mm pore size membrane, which is used to maintain a sterile root compartment and a non-sterile soil compartment. This is often used to determine the effects of root exudates on soil chemical properties or microbial biomass activity changes (Meharg and Killham, 1991; Yeates and Darrah, 1991). A sterile root compartment is maintained so that exudates are not degraded by microorganisms prior to entering the soil. The second, more common approach involves the use of a 30–60 mm mesh to separate non-sterile roots from non-sterile soil. This mesh size usually does not allow penetration by root hairs (diameter 50–100 mm) but is sufficient to allow passage of microorganisms. In addition to this, successive samples away from the rhizosphere can be determined with a resolution of up to 10 mm using a microtome approach as described by Yeates and Darrah (1991) and Joner et al. (1995). However, the technique is not without its limitations, which include: (i) interference of water and nutrient flow by the imposition of membranes; (ii) the exclusion of rootgrazing mesofauna; (iii) the exclusion of the rhizoplane and endorhizosphere; and (iv) the difficulty of spatial isolation of individual root sections required for spatial analysis. Despite these limitations, however, this type of exclusion technique has been successfully used in numerous rhizosphere investigations (Gahoonia et al., 1994; Marschner, 1995; Alphei et al., 1996).


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15.3 Methods for Rhizosphere Soil Chemistry Once rhizosphere and bulk (non-rhizosphere) soil samples have been obtained by the techniques outlined above, the chemical properties of each root zone can be compared by standard techniques. A typical analysis may include pH, major cations (ammonium, calcium, magnesium, potassium, sodium, aluminium), anions (nitrate, sulphate, phosphate), micronutrients (zinc, copper, iron) and dissolved organic carbon and nitrogen. Typically a variety of measures of nutrient status are performed including water or resin extraction, concentrated salt (KCl, NH4+-acetate), acid (HCl) or base (NaOH) extractions (Kowalenko and Yu, 1996; Trolove et al., 1996; Turrion et al., 1999; Zoysa et al., 1999). All of these provide information on differentially available fractions of soil nutrients (see Chapter 5). Soil solution can also be removed from larger rhizosphere samples using the centrifugal drainage method described by Jones and Edwards (1993). The details of the individual chemical analysis methods can be found in Weaver et al. (1994), Alef and Nannipieri (1995) and Sparks et al. (1996). There are now a variety of non-destructive in situ techniques for looking at the distribution of nutrient ions and biological activity in the rhizosphere. In the case of soil nutrient status, soil solution samplers have been designed to be able to sample gross rhizosphere solution. These are simply microversions of those used to sample bulk soil solution (see Section 7.2) and can be obtained in either ceramic, polysulphone or other organic materials (Göttlein et al., 1996; Farley and Fitter, 1999). Although these do allow an estimate of rhizosphere solution chemistry to be made, the sphere of sampling is often unknown, they only operate in moist soils (10 mM) (Bazzanella et al., 1998). This technique is by far the most powerful; however, it requires micromanipulators, capillary pulling equipment and good dissecting microscopes and is labour intensive. The overlaying of agar or filter papers impregnated with colorimetric indicators on to exposed roots has also been widely employed for determining rhizosphere nutrient dynamics. This technique is ideal for semiquantitative studies in which spatial dynamics are particularly important, for example when distinguishing effects of the root tip, root hair zone and root base. If the colorimetric indicators are non-rhizotoxic, then multiple exposures can be performed over time; however, in most cases only single time point measurements have been recorded. This

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technique has been used widely for determining the spatial localization of pH changes in the rhizosphere and especially in response to changes in nitrogen and phosphorus nutrition. Other applications involve the study of rhizosphere phosphatase activity, root ferric reductase activity, Mn4+ reduction, solubilization of inorganic phosphate and the release of organic acids (Dinkelaker et al., 1993a,b). Oxygen microelectrodes have also been developed which again are capable of non-destructively determining rhizosphere O2 concentration in a mesocosm-type system (Revsbech et al., 1999). The advantages and disadvantages are discussed in Armstrong (1994). In addition, an oxygensensing reporter strain of Pseudomonas fluorescens has also been used for monitoring the distribution of low-oxygen habitats in soil (Hojberg et al., 1999), as have CH4-sensitive biosensors (Damgaard et al., 1998).

15.4 Methods for Rhizosphere Biological Activity In recent years, great advances have been made in soil microbial ecology, allowing not just the biochemical characteristics of soils to be determined, such as enzyme activity, but also the size, activity and community structure of the rhizosphere microbial population. When investigating biological activity in the rhizosphere it is advisable to adopt a multifaceted approach. Typically this will involve a basic measure of soil microbial biomass (Beck et al., 1997) and microbial activity, usually basal or substrate-induced respiration (Jensen and Sorensen, 1994), alongside standard measures of key soil enzymes involved in carbon, nitrogen and phosphorus cycling such as protease, deaminase, cellulase and phosphatase activity (Alef and Nannipieri, 1995; Naseby et al., 1998). Although these techniques are straightforward and can be performed easily in most laboratories, measures of soil microbial community structure often require specialist equipment. At present a number of approaches can be taken, including the use of restriction fragment length polymorphism analysis (RFLP) (Chelius and Lepo, 1999), randomly amplified polymorphic DNA analysis (RAPD) (Redecker et al., 1999), fatty acid methyl ester analysis (FAMES) (Olsson and Persson, 1999; Siciliano and Germida, 1999), 16S rRNA amplification and temperature gradient gel electrophoresis polymerase chain reaction (TGGE-PCR) (Schwieger and Tebbe, 1998; Heuer et al., 1999; Smit et al., 1999), Biolog™ (Germida and Siciliano, 1998), enterobacterial repetitive intergenic consensus sequence PCR (ERIC-PCR) (DiGiovanni et al., 1999) and phospholipid fatty acid profiling (PLFA) (Griffiths et al., 1999). The most convenient of these techniques is Biolog™; however, it provides information only on culturable microorganisms and has therefore been criticized (Howard, 1997; Lawley and Bell, 1998).


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Although rhizosphere biological activity can be determined in destructively sampled rhizosphere soil, this typically provides little spatial or temporal resolution. This may be especially important considering that typically only 10% of the rhizoplane is colonized and that where colonization does occur it can be highly dependent on root architecture, such as cell junctions and the endorhizosphere. However, new techniques are now available which allow microbial activity and biomass of specific species to be determined in situ and in vivo. Further, the spatial and temporal dynamics can also be determined with high accuracy. The success of these techniques, however, relies on light emission from samples and therefore experimental design is critical to their success. For this reason, experiments are typically conducted in small microcosms and can only be performed in the laboratory. One of the main approaches has been to genetically modify target organisms with a bioluminescent luciferase (luxAB) gene, followed by the introduction of the tagged organism into microcosms. The amount of light produced by organisms established in the rhizosphere can subsequently be detected at a gross scale using commercial luminometers or with reasonable spatial resolution (0.05–10 cm) using a CCD-cooled camera (Deweger et al., 1991; Rattray et al., 1995; Prosser et al., 1996). The amount of light produced by the tagged organisms can be correlated with microbial activity, allowing target organism activity as a function of time and space to be determined (Eberl et al., 1997; Hagen et al., 1997; Wood et al., 1997). In addition, the luxAB gene insert can be placed into the target organism behind specific gene promoters, allowing the expression of individual microbial metabolic pathways to be assessed, such as those involved in nitrogen and phosphorus starvation (Kragelund et al., 1997; Jensen et al., 1998; Milcamps et al., 1998). Another similar approach has been the introduction of microorganisms tagged with a green fluorescent protein (GFP) into rhizosphere soil. This allows detection of introduced microorganisms using standard epifluorescence and confocal microscopes. The constitutively expressed GFP can be introduced as a quasi-stable tag, allowing total population numbers to be evaluated. In contrast, the introduction of an unstable GFP tag allows the expression of specific genes and metabolic pathways to be determined, thereby providing measures of microbial activity (Gage et al., 1996; Normander et al., 1999; Ravnskov et al., 1999; Tombolini et al., 1999). In a similar fashion to that of the luxAB genes, the GFP cassette can also be placed behind specific promotors, allowing the study of specific metabolic pathways such as those involved in nitrogen fixation (Egener et al., 1998).

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15.5 Quantification of Root Carbon Loss into the Rhizosphere The driving force for many rhizosphere processes is carbon loss from the root. Quantification of this carbon loss can be achieved in a variety of ways. The first involves whole-plant radiolabelling by continuous or pulse feeding the leaves with 14CO2 either in the laboratory or in the field, as described by Meharg and Killham (1988). Although this approach allows total plant carbon budgets to be calculated it does not provide information on the type of carbon compounds being released and has been criticized (Meharg, 1994). To determine the spectrum of exudate loss, a number of approaches can be taken, including extraction of rhizosphere soil solution by centrifugal drainage (Ström, 1997), the collection of charged exudate compounds by ion-exchange resins (Kamh et al., 1999) and the growth of plants under sterile conditions followed by collection in hydroponic or sand culture (Jones and Darrah, 1993; Hodge et al., 1998). The problem associated with determining carbon loss in sterile systems is discussed in Jones and Darrah (1993). The analysis of exudate components is usually carried out by standard gas chromatography, capillary electrophoresis, high-pressure liquid chromatography and enzymatic procedures, details of which can be found in any standard biochemical methods text. In addition, sites of carbon loss can also be determined semiquantitatively with bacterial biosensors (Jaeger et al., 1999).

15.6 Rhizosphere Mathematical Modelling Mechanistically based computer simulation models are available to predict the spatial and temporal dynamics of complex rhizosphere processes (see Fig. 15.1). They incorporate parameters such as diffusion and mass flow of nutrients and water, nutrient sorption/desorption and fixation reactions, soil solution chemical equilibria and characteristics of root nutrient influx, root radius, root hairs and shoot demand. Most of the models developed have been used for predicting nutrient uptake at the root, whole plant and field level (Sadana and Claassen, 1999). The construction of these models and examples of their output are extensively described in Tinker and Nye (1999), Cushman (1984) and Barber (1995). Other types of models have been developed from this basic design into which redox reactions have been incorporated (for rice-paddy soils) or for work in model rhizosphere systems (Jones and Darrah, 1993; Kirk and Solivas, 1994; Calba et al., 1999). The latest generation of models has been used to describe bacterial dynamics in the rhizosphere in response to root carbon additions (Darrah, 1991b; Scott et al., 1995), whereas others have fused nutrient and bacterial


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models to describe nutrient cycling involving biodegradable nutrientmobilizing organic ligands (Geelhoed et al., 1999; Kirk, 1999; Kirk et al., 1999). For a recent review see Toal et al. (2000).

Chapter 16 Soil Macrofauna P. LAVELLE,1 B. SENAPATI2

AND

E. BARROS3

1Laboratoire

d’Ecologie des Sols Tropicaux, 32 rue Henri Varagnat, 93143 Bondy Cedex, France; 2Ecology Section, School of Life Sciences, Sambalpur University, Jyoti Vihar, 768019, Orissa State, India; 3National Institute for Research in the Amazon (INPA), CP 478, 69011–970 Manaus, AM, Brazil

16.1 Synopsis There is growing awareness that the sustainability of agricultural systems will largely depend on the adequate management of biological resources (Woomer and Swift, 1994). In the suite of factors that determine soil fertility, soil organisms are proximate regulators of nutrient availability and soil structure (Lavelle, 1997). Their importance as a resource for agriculture has been underestimated because often only constraints imposed by climate and chemical parameters of soil fertility have been considered as aspects that could be influenced by management. Since the 1980s and the new soil fertility paradigms proposed by the Tropical Soil Biology and Fertility Programme (TSBF), emphasis has been placed on the cultivation of plant species that ensure nitrogen fixation and provide organic matter and nutrients for the system while generating sufficient income to sustain farmers’ livelihoods (Sanchez, 1994; Woomer and Swift, 1994). In addition to plant leaf litter and root systems, whose important roles in carbon and nutrient cycling, maintenance of soil structure and biological activity have been discussed in previous chapters of this book, soil invertebrates constitute another biological component that regulates basic soil processes such as soil aggregation, porosity and organic matter dynamics. Their management is now considered important, especially because most current land-use practices either destroy soil faunal communities or severely deplete their diversity, often with negative consequences for soil fertility (Chauvel et al., 1999; Lavelle et al., 1999). In this chapter, the composition and functional structure of soil © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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invertebrate communities are discussed, their effects on soil fertility are explained in broad terms, the effects of agroforestry practices on their communities are analysed, and techniques to assess their composition, abundance and activity are described.

Soil organisms and their functional domains Soil organisms have evolved in an environment that imposes three major requirements: (i) to move in a compact environment with a loosely connected porosity; (ii) to feed on low-quality resources; and (iii) to adapt to the occasional drying or flooding of the porous space (Lavelle, 1997). A continuum of adaptive strategies based on size is observed in soils, from microorganisms to macroinvertebrates (Swift et al., 1979). Soil invertebrates have been divided into micro-, meso- and macrofauna depending on the average size of the individuals. Microfauna comprises invertebrates 1 cm). In more general terms, a soil macrofauna taxon may be defined as an invertebrate group found within terrestrial soil samples which has more than 90% of its specimens in such samples visible to the naked eye (P. Eggleton et al., unpublished). According to this definition, large individual Collembola or Acari should not be considered as macrofauna. Microorganisms have developed the ability to digest every substrate present in soils. Unable to move, they depend on other organisms to transport them to where they can come into contact with the substrates on which they feed. Soil invertebrates have limited abilities to digest the complex organic substrates of the soil and litter, but many have developed interactions with microflora that allow them to exploit soil resources. With growing size, the relationship between microflora and fauna gradually shifts from predation to mutualisms of increasing efficiency. Excrement of invertebrates is of utmost importance in the evolution of organic matter (Martin and Marinissen, 1993), in the formation and maintenance of soil structure and, over long periods of time, in specific pedological processes referred to as zoological ripening of soils (Bal, 1982). Three major guilds of soil invertebrates, described below, may be distinguished on the basis of the relationships that they have with soil microorganisms and the kind of excrement and other biogenic structures that they produce. •

Micropredators mainly comprise microfauna that feed on bacteria, fungi and other micropredators. Microfauna do not seem to produce recognizable solid excrement and so their effect on soil organic matter

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•

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dynamics is not prolonged in structures that are stable for some time after deposition. They do, however, have a significant impact on the population dynamics of microorganisms and the release of nutrients immobilized in microbial biomass (Trofymow and Coleman, 1982; Clarholm, 1985). This process is especially developed in the rhizosphere (see Chapter 15). Predatory Acari or Collembola and even larger invertebrates (earthworms) may extend this food web over several trophic levels (Hunt, 1987; Moore et al., 1993). Litter transformers mainly comprise mesofauna and large arthropods that normally ingest purely organic material and develop an external (exhabitational sensu; Lewis, 1985) mutualism with microflora based on an external rumen type of digestion (Swift et al., 1979). Litter arthropods may digest part of the microbial biomass or develop mutualistic interactions in their faecal pellets. In these structures, organic resources which have been fragmented and moistened during the gut transit are actively digested by microflora. After some days of incubation, arthropods often re-ingest their pellets and absorb the assimilable organic compounds that have been released by microbial activity, and occasionally part of the microbial biomass itself (see, for example, Hassal and Rushton, 1982; Szlàvecz and Pobozsny, 1995). Finally, the ecosystem engineers comprise macrofauna, mainly earthworms, termites and ants, that are large enough to develop mutualistic relationships with microflora inside their proper gut. These organisms usually ingest a mixture of organic and mineral elements. Large faecal pellets (in the range of 0.1–2 cm and more) may be the component elements of macro-aggregate structures and participate prominently in the formation of stable structures through the regulation of porosity, aggregation, bulk density and surface features (Bal, 1982; Blanchart et al., 1999; Decaëns et al., 1999). These organisms also build large structures, such as mounds and networks of galleries and chambers, which have significant impacts on the evolution of soils over medium time scales. Ants may be considered part of this group, although the vast majority of them only use soil as a habitat and have a limited impact on soil organic matter dynamics.

Whenever conditions are suitable for their activities, macroinvertebrates, and especially earthworms and termites, become major regulators of microbial activities within their sphere of influence, referred to as the termitosphere of termites and the drilosphere of earthworms (Lavelle, 1997). Here, they also determine the abundance and activities of smaller groups of soil fauna (Dash et al., 1980; Yeates, 1981; Boyer, 1998; Loranger et al., 1998). These functional domains also include the rhizosphere, in which roots are the major determinants of microbial and faunal activity (see Chapter 15).

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Communities of organisms in soil are thus largely interactive, with a clear hierarchical organization dominated by ecosystem engineers. This justifies putting special emphasis on the management of macroinvertebrate communities for manipulating the whole biological activity of soils that they actually regulate inside their functional domains. The sampling of macroinvertebrate communities, which comprise most ecosystem engineers and litter transformers, is therefore proposed as a suitable proxy for overall soil biological activity (see Section 16.2). Attempts have been made to further classify macroinvertebrates into functional groups with a real functional meaning (Bouché, 1977; Grassé,

Box 16.1. Physical and chemical properties of biostructures deposited by soil invertebrate engineers at the soil surface in the eastern Llanos of Colombia. In a natural savanna of eastern Colombia, structures deposited at the soil surface by 17 species of soil invertebrate engineers were analysed for their chemical and physical characteristics and data were treated with principal component analysis (PCA) (Fig. 16.1). The analysis separated structures into three groups corresponding to large taxonomic units (earthworms, ants and termites). Earthworm casts were characterized by high values of bulk density, pH and contents of organic matter and cations. Termite structures differed from earthworm casts in having lower pH, lower bulk density and larger diameter, which reflects a greater structural stability. Ant structures had the highest aluminium saturation and the lowest amounts of soil organic matter and cations. This study confirms that physicochemical properties of biostructures are related to the faunal taxonomic units that produce them, suggesting that these represent, to a certain extent, functional groups. An exception was the group of Termitidae sp. 4, whose sheathings were similar to ant surface nest material (Decaëns et al., 2001). Fig. 16.1. (Opposite) Biostructures deposited on the soil surface of the eastern Llanos of Colombia by 17 species of soil invertebrate engineers as ordinated with principal component analysis (PCA) according to their physical and chemical properties: (a) Eigen values for extracted factors (the first two axes represent 59.6 and 22.6% of total inertia, respectively); (b) correlation circles associated with the first two axes; (c) ordination of biostructures in the plane delimited by Factors 1 and 2 of the PCA (small circles are means of points representing a category of structures produced). Abbreviations: %Corg, per cent organic carbon; PBrayII, available phosphorus content (BrayII); Ntotal, total nitrogen; Ptotal, total phosphorus; Al, exchangeable aluminium; Ca, exchangeable calcium; Mg, exchangeable magnesium; K, exchangeable potassium; %Al, per cent aluminium saturation; BD, bulk density; MWD, mean weighted diameter of aggregates; V, earthworm casts; T, termites (nests of all species except for surface sheathings of Termitidae sp. 4); A, ant surface nests (reproduced with permission from Decaëns et al., 2001).

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Axis 2

(b)

Al K Ptotal Mg %Corg PBrayII Ntotal

%Al

Axis 1 Ca (a)

MWD

pH BD Axis 2

(c)

Termitidae sp. 4 Atta sp.

Termitidae sp. 1 T

Camponotus sp. A

Nasutitermitinae sp. Termitidae sp. 3

Pheidole sp.

Acromyrmex landolti Trachymyrmex sp.

Axis 1

Martiodrilus carimaguensis

2.7 –2.7 6.8 –4.3

V

Andiodrilus yoparensis

1984). The effects of disturbances or changes in the environment on invertebrates indicate that large taxonomic units (such as termites and earthworms) are often fairly homogeneous groups in terms of response (sensu; Lavorel et al., 1997) and represent a rough approximation to real functional groups, that is, groups of species that perform similar functions within ecosystems (Steneck, 2001). For example, in natural savannas of

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Table 16.1. Functional categories of earthworms.a Ecological category

Habitat

Diet

Pigmentation

Epigeic Anecic

Litter Litter and soil

Litter feeder Feeds on litter and soil

Endogeic (polyhumic) Endogeic (mesohumic)

Topsoil

Endogeic (Oligohumic) aAfter

Composting ability

Galleries

Size

Very good Fair

Nil or very limited Long-lasting vertical

Small Medium to large

Rich soil feeder

Fully pigmented Anterodorsally pigmented Nil

Poor

Small

A horizon

Bulk Ai soil feeder

Nil

Nil

A and B horizons

Deep soil feeder

Nil

Nil

Horizontal, partly refilled with casts Extensive horizontal galleries refilled with casts Horizontal galleries refilled with casts

Medium

Large to very large

Bouché (1977) and Lavelle (1983). P. Lavelle et al.

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Colombia, the physical and chemical properties of biostructures produced by 17 different macroinvertebrate species were clearly related to large taxonomic units, with only one exception (Decaëns et al., 2001; see Box 16.1, p. 306). Within large taxa, species can be further subdivided into functional groups of species with distinct roles in the soil. Earthworms have been classified into five different categories depending on their feeding habits and location in the soil (Table 16.1; Lavelle, 1983). These categories have different distinct effects on soil function. Epigeics are useful agents in the production of compost but poor bioturbators (i.e. organisms that mix litter and soil). Anecics create persistent vertical below-ground networks of galleries that open at the soil surface; they also produce large numbers of casts, which they deposit in soil macropores or at the soil surface. Endogeics are the most active bioturbators; they deposit most of their casts in the soil and dig horizontal below-ground galleries that they refill with their casts. They have been further divided into poly-, meso- and oligohumics with decreasing average concentrations of organic matter in the ingested soil. Termites may be broadly classified on the basis of their feeding habits, general organization of their nests, and their location in the environment. Nests may be located in five main sites: (i) within the wood of living and dead trees and in fallen timber; (ii) subterranean nests; (iii) epigeal nests on the soil surface; (iv) arboreal nests hanging from branches or stuck to tree trunks; and (v) within the nests of other termite species (Noirot, 1970). Termites may also be separated into four groups on the basis of their feeding habits: (i) wood feeders; (ii) fungus cultivators; (iii) grass harvesters and surface litter feeders; and (iv) humivorous termites (Josens, 1983). A small number of species also regularly attack living plants. Wood feeders may be further separated into several groups, such as dry-wood, wet-wood or arboreal termites (Abe, 1987). No specific classifications have been proposed so far for other soil and litter macroinvertebrates. A classification of ants based on their effects on bioturbation and transfers of organic matter in soil is clearly needed. Litter transformers (mainly Diplopoda, Isopoda and detritivore Coleoptera) seem to be a rather homogeneous functional group, although they may differ significantly in their response to environmental factors. Predators are mostly found in Myriapoda (Chilopoda), Insecta (Coleoptera and ants) and Arachnida. Effects of macroinvertebrates on soil processes Fauna effects on soil physical properties Soil invertebrates influence soil processes directly by their digestion processes and the formation of biogenic structures. The latter effect is by

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far the most important since it determines basic soil physical properties and further steps of organic matter dynamics. Soil aggregation in most humid tropical environments is determined by the activity and functional diversity of the main ecosystem engineers. In moist savannas of Côte d’Ivoire, earthworms may produce up to 1200 t of casts ha–1 year–1, of which only 30 t (2.5%) are deposited on the soil surface (Lavelle, 1975). The earthworm community comprises compacting species that ingest small aggregates and produce large compact structures, the accumulation of which tends to increase the bulk density of the soil (Blanchart et al., 1999), and decompacting species, which have opposite effects through the fragmentation of large aggregates into smaller ones and/or the creation of pores. The combined effects of these two broad functional categories maintain soil physical and hydraulic properties in a highly dynamic state. Ants and termites also have strong impacts on soil physical parameters (Lobry de Bruyn and Conacher, 1990). Some Macrotermitinae may open access holes at the surface, which they use for foraging at night. The overall surface of these openings has been estimated at 2–4 m2 ha–1 in a dry savanna of Kenya (Lepage, 1981). Gallery lengths of up to 7.5 km ha–1 have been estimated for soils associated with the mounds of Macrotermes michaelseni (Darlington, 1982; MacKay et al., 1985; MacKay and Whitford, 1988). In the soils of Darlington’s (1982) study area there was also the equivalent of 90,000 storage chambers per hectare. It can be calculated that the voids formed by termite galleries and related structures occupied approximately 0.4% of the soil volume to 20 cm depth at this site (Lavelle and Spain, 2001). The effect of soil fauna on soil structure can be surprisingly rapid. Barros et al. (2001) showed that the structure of an Amazonian Oxisol could be drastically modified by changes in faunal activity in only 1 year. Soil monoliths taken from a pasture soil that had been severely compacted by an invasive earthworm were transported to a nearby forest. After 1 year of exposure to the diverse fauna of the forest, soil of the exposed monoliths had recovered the overall porosity and the percentage of macroaggregates had returned to the situation found in forest soil. While this example demonstrates the importance of decompacting fauna species for the maintenance of soil hydraulic properties, compacting species may also play an important role through the stabilizing effect of the large and dense aggregates that they produce on soil structure which protects against mechanical stress and maintains organic matter in the interior of these dense aggregates (see below and Martin, 1991). Furthermore, an earthworm species that compacts the superficial soil horizons may increase soil porosity in the subsoil (Barros et al., 2001). The dynamic equilibrium between these antagonistic activities has a strong influence on soil physical, chemical and biological properties, and illustrates the importance of the functional diversity of soil organisms for soil functioning and fertility.

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Biogenic structures produced by ecosystem engineers are often rather resistant, especially after having experienced a few wetting and drying cycles. This is the case for earthworm casts that are highly unstable when fresh and soon acquire high stability after drying (Shipitalo and Protz, 1988; Blanchart et al., 1993; Hindell et al., 1997). Dry surface casts of Martiodrilus carimaguensis, a large anecic earthworm of the Colombian Llanos Orientales, can resist heavy rainfall and persist for several months before rainfall and runoff break them down into stable aggregates that form a mulch on the soil surface (Decaëns, 2000). Unless they are destroyed by another invertebrate, biogenic structures created inside the soil may persist for long periods of time after the organisms that produced them have died. Process regulation by ecosystem engineers may therefore persist long after they have disappeared, and degradation of soil properties due to their elimination by cropping practices, or any other disturbance, is often ignored because the link between the organism and the soil properties is not evident. Soil texture in the upper soil horizons may also be modified by macroinvertebrate activities as a result of the regular deposition of finely textured biogenic structures at the surface. This process has been hypothesized to generate stone-lines found at some depth in many tropical soils (Williams, 1968; Wielemaker, 1984; Johnson, 1990). In other circumstances, fine particles deposited by fauna at the soil surface may be washed away by runoff and leave a coarse-textured surface horizon (see Chapter 17). Fauna effects on soil organic matter dynamics The effects of soil invertebrates on soil organic matter dynamics are disproportionate to their size and numbers. Direct mineralization of soil organic matter by earthworms was estimated at 1.2 t ha–1 year–1 in a natural savanna of the Côte d’Ivoire (Lavelle, 1978). This corresponds to approximately 10% of all carbon mineralized in this soil, and the overall contribution of all soil fauna to carbon mineralization at this site probably does not exceed 20% (Lamotte, 1989). Indirect effects on organic matter dynamics are much larger, especially as they extend over several time scales. At a short time scale of days to weeks, the organic matter that has been ingested by invertebrates and egested in their faecal pellets undergoes further mineralization caused by a strong enhancement of microbial activity, labelled the ‘Sleeping Beauty effect’ by Lavelle (1997). At a longer time scale of months to years, organic matter occluded in compact biostructures like earthworm casts or walls of termite mounds may be physically protected from decomposition. Therefore, invertebrates that stimulate mineralization of organic matter at small scales of time and space tend to protect organic matter from further decomposition at much

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larger scales. The coexistence of these two processes interacting at distinct scales may lead to misinterpretation of some experimental results. Elimination of invertebrate populations only suppresses short-term effects, whereas long-term protection of organic matter in biogenic structures will last as long as these structures are maintained. However, Blanchart et al. (2003) have shown that this regulation of soil organic matter dynamics via the effects on soil physical structure was only significant in soils with low clay contents and soils with inactive clay minerals of the 1:1 type, but not in soils with smectites and other 2:1 clay minerals.

Effects of agroforestry practices on soil macrofauna Hierarchical determinants of soil function in agroforestry systems Soil function is controlled by a suite of hierarchically organized determinants, the relative importance of which depends on the scale at which they operate (Lavelle and Spain, 2001). Agroforestry practices will affect soil function as they act on these determinants. At the top of the hierarchy are climatic factors such as moisture and temperature regimes. At the next level down are edaphic parameters with special importance accorded to the nature and abundance of clay minerals and the overall nutrient richness of soil, which is partly determined by the nature of the original bedrock. At the next level down comes the quality of organic matter produced and other parameters linked to the plant communities that are present. Macroinvertebrate communities are dependent on these determinants and constraints imposed by biogeographical factors, such as the local history and spatial distribution of land-use systems, which impinge on the maintenance of diversity and colonization of newly created cropping systems from adjacent areas. These communities, in turn, are themselves a proximate determining factor for microbial activities in the soil. The few available data sets reflect this diversity of determinants of soil invertebrate communities (Lavelle and Pashanasi, 1989; Lavelle et al., 1994; Loranger, 1999). Agroforestry systems appear to have rather specific fauna communities compared with conventional cropping systems. Trees in agroforestry systems affect living conditions of invertebrates by their shade, their deep and perennial root systems, and the quantity and quality of their litter (Tian et al., 1995a; Vohland and Schroth, 1999), and by preventing soil tillage in at least part of the system. Tillage has a significant negative impact on soil macroinvertebrates. There is a direct effect through disruption of the soil structure and the destruction of either the invertebrates themselves or the structures that they inhabit. In the longer term, a combination of microclimatic and trophic

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effects prevents populations from building up rapidly since soil is left bare for some time after each tillage (House, 1985; Haines and Uren, 1990). Soil microclimate is affected by agroforestry practices in two different ways. Soil under trees is covered with leaf litter and shaded, so that the upper centimetres of soil are protected from overheating and drying (Lal, 1986). On the other hand, the perennial nature of trees, their often high evapotranspiration and the rather deep distribution of roots may cause more intensive and deeper drying of the soil during the dry season than in areas with annual crops or other herbaceous vegetation, and this may affect soil fauna. The quality and abundance of organic inputs is a key feature in the maintenance of diverse soil macroinvertebrate communities (Lavelle et al., 2001). The provision of habitats and food for litter invertebrates favours the maintenance of specific communities of epigeic invertebrates (mainly arthropods such as Myriapoda, Coleoptera, ants or surface termites, and epigeic earthworms) and anecic invertebrates which dwell in the soil but feed, at least partly, on surface leaf litter, such as termites and earthworms. Barros et al. (1999) showed an effect of litter cover on soil fauna in a comparison of two different agroforestry systems and three 10-year fallows in central Amazonia. In a system with still young trees, litter cover was low and the macroinvertebrate community had a lower overall abundance and diversity and tended to live deeper in the soil than in the other sites with

Fig. 16.2. Relationships between litter dry matter, faunal density (a) and faunal biomass (b) for different sampling positions in a multistrata agroforestry system with Brazil nut trees (Bertholletia excelsa), cupuaçu (Theobroma grandiflorum), annatto (Bixa orellana), peach palm (Bactris gasipaes), a Pueraria phaseoloides cover crop and spontaneous grass as well as a peach palm monoculture in central Amazonia (means and s.e.; reproduced with permission from Vohland and Schroth, 1999).

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older trees and more abundant litter. In the same region, Vohland and Schroth (1999) observed a close relationship between the amount of litter and the abundance of litter-dwelling macroinvertebrates. They estimated the amount of litter necessary for maintenance of an active fauna in the litter system at 3–6 t ha–1 (Fig. 16.2). The amount of soil organic matter which is assimilated by an earthworm community in a natural savanna of Côte d’Ivoire has been estimated at 1.2 t ha–1 year–1 (Lavelle, 1978). Based on this number, one could assume that if agroforestry practices succeeded in supplying 2 t ha–1 year–1 of assimilable organic matter this should be enough to sustain macrofaunal activity at a suitable level. We do not know, however, how much litter of what quality must be applied to the soil surface to sustain a litter biomass of 3–6 t ha–1 as a habitat for litter-dwelling invertebrates while providing 2 t ha–1 year–1 of organic matter for digestive assimilation. This question needs to be resolved for various agroforestry practices on different sites, taking the quantity and quality of available organic materials into consideration. Assimilability of plant residues is greatly determined by their chemical composition, especially lignin, polyphenol and nitrogen contents (see Chapter 6), and this influences the composition of the faunal community. In general, the ratio of termite to earthworm abundance is dependent on the quality of the organic residues; termites tend to predominate when low-quality organic substrates are available whereas earthworms thrive better with high-quality residues. In rubber plantations of southern Côte d’Ivoire, for example, xylophagous and other termites dominate in the early phases when trunks and branches of the original forest are left on the soil surface. At later stages, 10–20 years after plantation establishment, most woody litter has been eaten up and transformed by termites, and endogeic earthworms, which presumably feed on termite excreta, become dominant (Gilot et al., 1995). Tian et al. (1993) showed similar effects with different mulch covers in agroforestry systems in Nigeria. In silvopastoral systems, the use of improved pastures with selected African grasses and legumes, as well as grazing, generally increases earthworm abundance and the ratio of earthworm to termite population densities (Decaëns et al., 1994; B. Senapati, unpublished data). Cover crops and grass fallows may have similar impacts depending on the quality of organic matter produced. In a survey of soil macroinvertebrate communities in the Brazilian Amazon, E. Barros et al. (unpublished) found that mean values of the termite:earthworm density index, calculated from population densities, ranged from 0.2 in pastures to 7.9 and 8.8 in secondary forests and agroforestry systems and 20.4 and 21.4, respectively, in fallows and crops. The similarity between fallows and crops in this case indicates that the fallows were still young and so there was relatively little production of leaf and woody litter by the trees. Vohland and Schroth

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(1999) found that in Amazonian agroecosystems the fleshy harvest residues of peach palm (Bactris gasipaes), grown for heart-of-palm production, harboured diplopods and other fauna groups sensitive to desiccation, which, in the primary forest of the region, are typically found in rotting stems of fallen trees. Tree species effects within agroforestry systems The distribution and nature of plants in an agroforestry system will greatly influence macroinvertebrate communities. The particular trophic and microclimatic conditions in the proximity of trees may significantly shape the invertebrate community and lead to single-tree effects on their distribution (Blair et al., 1990; Boettcher and Kalisz, 1991). In a rainforest of French Guiana, endogeic earthworms were concentrated below the litter of an unidentified tree species of the Qualea genus but were totally absent from litter of another tree, Dicorynia guianensis, only 20 m away (Wardle and Lavelle, 1997). Distribution patterns of litter fauna related to the presence of individual trees have also been observed in a multistrata system with four tree crop species, a leguminous cover crop and spontaneous grasses in central Amazonia (Vohland and Schroth, 1999). Some fauna groups showed specific preferences for certain litter types. For example, earthworms were most abundant in the litter of the cover crop, Pueraria phaseoloides, peach palm (Bactris gasipaes) and annatto (Bixa orellana), but were absent from the hard and rather dry litter of cupuaçu (Theobroma grandiflorum) and Brazil nut (Bertholletia excelsa). Similar preferences were shown by millipedes and beetles (Fig. 16.2). In another multistrata system established after pasture in the same region, Barros et al. (1999) found a dominance of mesohumic endogeic earthworm species in the soil under cupuaçu and Brazil nut trees and of polyhumic earthworm species under mahogany (Swietenia macrophylla) and passion fruit (Passiflora edulis) within the same system. Presumably, this difference also reflected the low litter quality of cupuaçu and Brazil nut and, for young cupuaçu trees, the small quantity of litter produced. At a landscape scale the distance from an agroforestry plot to areas of natural vegetation (where it still exists) and the influence of neighbouring types of land use are expected to affect the composition of fauna communities. In central Amazonia, the colonization of agroforestry plots by a large anecic glossoscolecid earthworm seemed to depend on contact with natural forest (E. Barros, unpublished). However, adequate data sets to properly test this hypothesis are lacking. The role of biodiversity Agroforestry systems are generally considered to have a positive effect on the conservation of biodiversity compared with simpler agricultural

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systems (Fragoso and Rojas, 1994; Griffith, 2000). This is especially the case when their structure is close to that of the original ecosystem (Decaëns, 2000). Conservation of biodiversity is important both from a general ethical point of view and because it may be a precondition for the sustainable use of soils. However, this latter point is still poorly understood and limited to a few hypotheses (Hooper et al., 2000; Lavelle, 2000). It is assumed that below-ground faunal diversity is influenced by the diversity of plants growing at a site and vice versa. However, the relationship is complex since a large number of interactive processes are involved. Little is known about the role of diversity in soils under field conditions beyond experimental evidence that a minimum diversity directly influences rates of basic ecosystem processes such as mineralization of carbon and nutrients (Mikola and Setala, 1998). The relationship between species richness and functional diversity seems to be rather loose since some groups, like spiders or Coleoptera, may be locally represented by more than 100 species with high apparent functional redundancy whereas termites would only have 20–40 species and earthworms approximately ten species. The latter two groups may actually occupy much wider niche spaces than non-social arthropods because of their social organization and/or ecological plasticity (Lavelle and Spain, 2001). In some cases, significant degradation of soils has been related to an imbalance between functional groups, such as invasion of an Amazonian pasture by a peregrine earthworm species, which accumulated large amounts of compact structures close to the soil surface and, in combination with depletion of decompacting species (ants and termites), led to the formation of a continuous surface crust, which impeded plant growth (see above and Chauvel et al., 1999). The density of the compacting earthworm species was much lower in a small woody area (E. Barros and T. Decaëns, unpublished). The diversity of food sources and habitats characteristic of agroforestry systems may confer a lower susceptibility to such ‘biodiversity accidents’ than is likely in less diverse agricultural contexts. Specifically, the presence of leaf and woody litter at the soil surface is likely to stimulate the activity of decompacting species such as ants and termites, with positive effects on water infiltration (Mando and Miedema, 1997). Conclusions and research needs Agroforestry systems are a promising alternative to the intensive monocropping systems which were promulgated during the 40 years or so of the green revolution. Soil macroinvertebrates, especially the group referred to as ecosystem engineers, are part of the biological resources that need to be managed in these systems. Invertebrates are not only indicators of the health of the system; they are also actors participating in the maintenance of soil physical properties, short-term recycling of nutrients

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and longer-term protection of nutrients and organic matter in their biostructures. The direct management of soil invertebrate communities, by, for example, inoculation with earthworms, is difficult and costly. This makes understanding the effects of plant communities and management practices on invertebrate communities very important, so that these can be optimized through appropriate design and management of agroforestry and other land-use systems (Senapati et al., 1999; Lavelle and Spain, 2001). In addition to the maintenance of a favourable microclimate and the avoidance of tillage in at least part of the system, the greatest potential of agroforestry practices to benefit soil invertebrates lies in the provision of diverse and abundant organic resources. More research, however, is needed into ways of optimizing the supply of organic resources in different agroforestry practices and under various pedoclimatic conditions, with special attention given to the effects of litters of different quality, including mixtures of different litter types, on invertebrate communities. Generating information to reconcile the partly contradictory objectives, on the one hand, to provide sufficient assimilable organic matter (through rapid litter transformation) and, on the other hand, to maintain a continuous litter layer that protects the soil and serves as a habitat, requires an integrated research approach using experimental and modelling techniques. The composition of communities is also a relevant issue to address since the lack of functional diversity may have unexpected negative effects on soil function (Chauvel et al., 1999). The composition of an invertebrate community depends on a suite of determinants that operate at different scales of time and space. Geographical constraints determine regionally or locally the pool of species present. Distance from a plot to reserves of species, such as primary or secondary forest, and the size and shape of the plot are significant primary determinants of the community composition. Inside the plot, the quality and quantity of litter produced by single plant components and the array of plant species present are of great significance. Sampling protocols and procedures for data analysis that are adapted to these sources of variability are necessary. They should integrate the spatial structure of the system at scales from landscape (watershed) to farming system and plot. The functional classification of invertebrates is another key issue to address. Functional classification systems are still in their infancy and rarely address the real effect of invertebrates on ecosystem function. Furthermore, no comprehensive classification for all soil invertebrates really exists. Future research should seek to interpret community compositions in terms of their effect on decomposition rates, organic matter dynamics over different time scales and effects on soil physical properties.

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Finally, the energy balance of faunal activities has to be considered to design appropriate management options. These should be designed to provide enough energy to sustain an adequate level of faunal activity while allowing sufficient storage of recalcitrant or physically protected organic matter for the maintenance of exchange characteristics, storage of nutrients and stabilization of soil aggregates (Lavelle et al., 2001).

16.2 Sampling of Macrofauna: the TSBF Methodology Sampling designs A method for the collection of soil macrofauna in the field, which had originally been proposed by Lavelle (1988), has been adapted to the needs of the TSBF (Anderson and Ingram, 1993) and has recently been reevaluated and improved within the Macrofauna Programme of the International Biodiversity Observation Year (IBOY; D. Bignell et al., unpublished). The original sampling design comprised ten samples of 25 cm × 25 cm and 30 cm depth, distributed every 5 m along a transect chosen on a random basis. A problem with this design has been the strongly aggregative distribution of most invertebrate groups in the field, which results in high variance and creates autocorrelation among the sampling points. To avoid this problem, it is now recommended that the distance between sampling points is increased, whenever possible, to at least 20 m. In agroforestry systems, two alternative protocols may be applied, which are stratified sampling or systematic sampling on a regular grid (see Section 3.4). With stratified sampling, samples of soil macrofauna are collected beneath trees of different species, if tree species effects are expected, and in the different individual components such as grass bands, crop strips or hedgerows. Ten samples per unit and certainly not fewer than five are recommended to detect significant differences between positions. When considering communities at the scale of a farm or watershed, sampling on a regular grid is recommended. This design provides an average estimate of soil macrofauna communities in large landscape units covered by the grid and allows a stratification a posteriori by separating points on the basis of the local plant cover. Another advantage of this protocol is that the spatial structure of the plot is explicit and can be analysed with geostatistical methods (see Section 3.6). This is of great value in landscapes with land cover mosaics where movements of fauna between adjacent plots may determine the composition of communities. Spacing between sampling points will depend on the size of the area and the number of samples that can be processed with the available resources.

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Hand-sorting The sampling area is delimited, preferably with a metallic frame corresponding to the size of the sample (25 cm × 25 cm) with vertical blades of 5–10 cm on all sides that help to fix the device in position and to cut through the litter and topsoil. If litter is present, it is collected and kept in a hermetic plastic bag for further sorting. The soil sample is then isolated by cutting the soil vertically to a depth of 30 cm with a flat spade or machete. A ditch is dug on one of the four sides, and successive 10 cm strata are cut and kept in separate plastic bags for further processing. Hand-sorting is carried out on large flat trays (40 cm × 60 cm and 5 cm deep). A handful of litter or soil at a time is spread in the tray to allow the capture of all individuals that can be seen with the naked eye. Controls of the accuracy of the method have shown that a variable proportion of individuals is found. This proportion may vary from 10% to almost 100% depending on the size, colour and mobility of the invertebrates (Lavelle, 1978; Lavelle and Kohlmann, 1984). In spite of the relatively low and variable efficiency of the sorting, the method has proven robust and highly repeatable in a wide range of tropical and temperate areas with forest, crops or grass cover (Lavelle and Pashanasi, 1989; Dangerfield, 1990; Decaëns et al., 1994; Lavelle et al., 1994). The results are normally used directly in multivariate analyses for the comparison of different sites or treatments. Where absolute values are important, rather than relative differences in community structure between sites, correction factors may be established for different faunal groups at the beginning of a sampling programme. This is done by sorting samples both in the field and by heat extraction (Berlese funnels) for litter and washing–sieving extraction for mineral soil (see, for example, Lavelle and Kohlmann, 1984). This approach could be especially useful in long-term studies at a single site. It will normally take one person-day to cut and hand-sort two or three samples in the field. Since variance is always very high due to the inherently heterogeneous distribution of soil invertebrates, the recommended number of samples does not guarantee that statistically significant differences between treatments or sampling positions are obtained, even if effects are relatively large.

Identification and data storage Although no precise guidelines on identification can be given at this stage, the invertebrates collected should be separated into a minimum list of major taxa for entry into a standardized database (Table 16.2; Fragoso et al., 1999). Whenever possible, more precise identifications should be used. Families will generally yield 30–60 entries. Identifications at the species

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Table 16.2. Minimum list of taxonomic units for soil macrofauna for inclusion in the MACROFAUNA database.a Earthworms Ants Termites Coleoptera adult Coleoptera larvae Total Coleoptera Arachnida Diplopoda Chilopoda Total Myriapoda Isopoda Dictyoptera (Blattoidea) Hemiptera Dermaptera Lepidoptera larvae Orthoptera Gasteropoda Other macrofauna aAfter

Fragoso et al. (1999).

level should be attempted if the requisite taxonomic expertise is available. General identification keys are not yet available but identification tools will be progressively released as part of the IBOY programme on macrofauna described above. These tools will be made available along with access to the MACROFAUNA database on the Internet (www.bondy.ird.fr/bondy.lest). The use of morphospecies or recognizable taxonomic units (RTUs) is recommended when identification is not possible (Williams and Gaston, 1994; Stork, 1995). Populations of each taxon are characterized by their numbers (individuals per m2) and biomass. Biomass is preferably expressed as fresh weight of preserved invertebrates. Expressing biomass as dry weight considerably increases the amount of work and does not increase comparability of results unless guts are emptied of their soil or litter contents.

16.3 Other Sampling Methods Depending on the purpose of the study and the available manpower, alternative methods may be used such as the rapid biodiversity assessment (RBA) method (Oliver and Beattie, 1993; Jones and Eggleton, 2000), or

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pitfall traps designed to catch the fauna that moves across the soil surface. Collection of biogenic structures deposited at the soil surface may be an acceptable surrogate to the evaluation of abundance, diversity and activity of invertebrate engineers.

Pitfall traps Pitfall traps are pots (approximately 10 cm in diameter and 15 cm in depth) that are fitted into the soil in order to catch the vagile fauna. Ants and non-social litter arthropods (Arachnida, Coleoptera, Myriapoda) are the most commonly caught invertebrates; earthworms, termites and all truly endogeic soil fauna are less efficiently sampled or missed altogether. The following protocol was designed to sample surface arthropods in 1 km2 square plots in European landscapes as part of the BIOASSESS programme (J. Niemela, unpublished). Sixteen sampling points are regularly distributed across the area. At each point, four traps are installed at the four corners of a 5 m × 5 m square. Traps 8 cm in diameter are partly filled with either ethylene glycol or propylene glycol. They are covered with a rain cover placed a few centimetres above the trap. The trapping period extends over a minimum of 10 weeks and traps must be checked frequently to avoid decay and loss of material. Other sampling designs are needed for studying small-scale patterns within agroforestry plots, such as stratified sampling under different tree species, or sampling shaded and unshaded or mulched and unmulched areas. Traps are particularly suitable for such purposes (Lavelle and Kohlmann, 1984; Hanne, 2001). Pitfall traps may be made more efficient or selective by adding specific food substrates such as jam and canned tuna fish (ants), vertebrate dung or faeces (scarabeid beetles) or wood pieces (xylophagous termites). A different trap design specifically for termites consists of plastic tubes of about 5 cm diameter with lateral openings which are inserted in the soil so that the openings are below the soil surface. The tubes are covered with a lid. Pieces of wood are placed inside the tubes and are regularly controlled for the presence of termites (Su and Scheffrahn, 1996). Preliminary tests with different types of wood are recommended (Hanne, 2001).

Collection of biogenic structures Soil ecosystem engineers produce a great variety of biostructures that they may deposit at the soil surface, hang on tree trunks or incorporate into the soil. Earthworms produce casts, middens, galleries and resting

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chambers. Earthworm casts are all made of faecal material that is generally richer in fine and organic particles than the bulk soil. Termites build epigeic nests of highly variable shapes and sizes, at the soil surface and/or in the soil, on tree branches or inside rotten trunks. They also dig a wide range of galleries and chambers and deposit surface covers and runways. Termite biofabrics and deposits may be made either of faecal material or material simply separated and worked with their mandibles. Non-faecal material may be either soil or carton, which is a soft mixture of excreta, undigested plant material and inorganic elements. The texture and organic contents of termite structures may be highly variable. Ants do not comprise geophagous populations and all their constructs are made with soil that has been removed with mandibles and deposited outside the endogeic nests, most often as accumulation of loose soil aggregates. In some cases, however, ants may build rather compact epigeic earthen nests, like Camponotus punctulatus in subtropical South America. Quantifying the abundance, diversity and composition of these structures may be considered as a functional substitute for the quantitative evaluation of soil macroinvertebrate communities (see Box 16.1). As such, this approach may be used to assess the impact of different agricultural practices. Sampling protocols need to be adapted to the local conditions. Large, long-lasting structures should be counted once or twice per year over large areas, whereas short-lived structures like most earthworm casts or ant deposits can be assessed by removing them from a surface and then measuring how many new structures are produced in a given period of time. This must be done in periods without rainfall, which may wash away the most fragile structures. Biostructures in the soil can be studied and quantified with soil micromorphological techniques (see Section 10.6).

16.4 Manipulative Experiments The effect of invertebrates on soil processes can also be assessed with manipulative experiments in the field. Litterbags are widely used to assess decomposition of litter under field conditions (see Section 6.3 and Anderson and Ingram, 1993). Litter is enclosed in a mesh made of an undecomposable material and left on or in the soil under field climatic conditions and exposed to the community of decomposers. The participation of invertebrates in decomposition is classically evaluated by using different mesh sizes that permit access by invertebrates of different sizes (e.g. 0.2, 2 and 5 mm). This technique has been widely used, so that it produces data that are readily comparable with other data sets. The major drawbacks are the confinement of organisms due to the effect of the mesh, an altered microclimate in the mesh bag and an imperfect contact

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with underlying litter or soil layers. For a detailed discussion of the method see Section 6.3. Effects of soil fauna and other biological and non-biological agents influencing the soil structure, such as roots or wetting–drying cycles, can be assessed by exposing undisturbed soil monoliths in a given soil environment and monitoring changes in their physical structure and other properties (Blanchart, 1992; Barros et al., 2001). The formation of large aggregates from a soil previously sieved at 2 mm under the influence of earthworms has also been studied with this method. Re-aggregation of soil by large compacting earthworms was quite fast, with the original level of aggregation being obtained after only 1 year (Blanchart, 1992).

Chapter 17 Soil Erosion M.A. MCDONALD,1 A. LAWRENCE2

AND

P.K. SHRESTHA3

1School

of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK; 2Environmental Change Institute, University of Oxford, 5 South Parks Road, Oxford OX1 3UB, UK; 3Local Initiatives for Biodiversity, Research and Development (LI-BIRD), PO Box 324, Bastolathar, Mahendrapool, Pokhara, Nepal

17.1 Synopsis There are two basic approaches to the study of erosion rates. In the first, sediment transport rates are monitored at the outlet from a catchment. Such measurements are relatively easy to make and they integrate the effects of erosion over a variety of land uses but there are problems in determining the source of the sediment within the catchment. The second approach involves measuring processes at a number of sampling sites within a catchment. The techniques vary with the processes involved. Such data are more difficult to collect but have obvious advantages in providing information about the spatial distribution of erosion rates within a catchment and the relative contribution of different land uses. The latter approach is the most applicable to agroforestry research at a field scale, seeking to investigate the effects of incorporating a tree component into a farmed environment. If both approaches are combined within the same catchment, the results of the former may serve as a test for the successful scaling up of the results of the latter. The need to evaluate rates of soil erosion in agroforestry arises from investigations on sloping land where soil transport is a major issue. The presence of trees in hillside landscapes complicates processes that affect soil erosion so that methodologies commonly used to evaluate it require adaptation for use in agroforestry, and it is important to have a precise definition of the objectives of the research. Considerable interest in agroforestry research has focused on the potential for erosion control, and the role of trees in the redistribution of soil and associated nutrients and moisture on farmed slopes. © CAB International 2003. Trees, Crops and Soil Fertility (eds G. Schroth and F.L. Sinclair)

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Trees in agroforestry systems can affect rates of erosion in several ways: (i) they can act as a semipermeable barrier which slows the velocity of surface runoff, and hence its erosivity; (ii) they can increase infiltration by the same mechanism and also through the improvement of soil structure (see Chapter 10) and hence reduce runoff volume (which reduces nutrient losses in runoff but can increase nutrient leaching; Ramakrishnan, 1990); (iii) they can protect the soil from the impact of rainfall by production of a litter layer and/or prunings that are used as a mulch; and (iv) they can act as a sieve barrier to eroding soil and hence change the particle size composition of eroded sediments. The presence of a tree canopy has contradictory effects on erosion. On the one hand, trees reduce the quantity of water that reaches the soil through crown interception (see Section 11.1). On the other hand, the erosivity of rainfall may increase because the size and spatial distribution of raindrops are modified during crown passage (Armstrong and Mitchell, 1988). Raindrops intercepted by tree leaves may coalesce on leaf surfaces before emerging as larger throughfall droplets, which are more erosive, beneath the tree crown. The distribution of drop sizes under trees with different leaf morphologies has been found to vary significantly, with larger leaves generally producing larger drop sizes (Calder, 2001), although drop size also depends on leaf surface characteristics and leaf inclination. The proportion of raindrops that are modified as opposed to passing through the canopy unaffected depends on crown cover and density. Tree height may also increase the kinetic energy and hence the erosivity of drops reaching the soil. Drops reach 95% of their terminal velocity in a free fall of 8 m (Young, 1997), so a further increase in height does not greatly increase their velocity when reaching the soil. Increased erosivity of throughfall over rainfall in agroforestry systems is often well understood by farmers (Thapa et al., 1995) and has been well documented scientifically (Wiersum, 1984). Some trees also produce high stemflow rates, which may increase erosion under certain conditions (Wiersum, 1985). In light of this, it is unsurprising that experimental work has clearly shown that it is not the presence of the tree canopies, but rather the direct soil protection offered by a continuous litter layer that is responsible for low erosion rates in undisturbed forest (Wiersum, 1985). This has clear consequences for erosion control in agroforestry systems, where the litter layer may be disturbed during crop cultivation. There are two principal ways of integrating trees and shrubs to control erosion in agroforestry, which differ in the relative importance of barrier as opposed to soil cover effects. These are: • •

dense, mixed agroforestry systems (multistrata systems and perennial tree crop combinations); and spatially zoned systems (contour hedgerows and other systems of trees and shrubs in contour-aligned belts).

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The former control erosion by means of a dense, surface ground cover of plants, litter or mulch, which, if maintained, is clearly effective (Wiersum, 1984; Young, 1997), though few experimental studies have been performed. Spatially zoned systems have received greater attention, and experimental results have been instructive in establishing the role of trees on slopes in increasing spatial heterogeneity in a number of soil properties (see, for example, the discussion of terrace formation below). In an agroforestry context, a fundamental question is how landscapes with trees are positioned in a continuum from pure cropland to natural forest (van Noordwijk et al., 1998) and it is important that the processes elucidated in studies of trees at various densities and in various spatial arrangements are considered in their landscape context.

Approaches to measuring soil erosion in agroforestry research Understanding the interaction of trees and slopes on the redistribution of soil and nutrients within the area being evaluated is of fundamental importance. Strong evidence has been found for a scouring effect in contour hedgerow systems on hill slopes whereby soil is lost from the upper part of the alley between hedgerows and accumulates in its lower part, leading to a transfer of surface soil properties between the two parts of the alley (Garrity, 1996). Usually, the effects of this redistribution of soil within the alleys is such that, after some time, the differences in soil properties between the upslope and the downslope side of an alley are greater than those between areas with and without hedgerows (Garrity, 1996; Agus et al., 1997; Poudel et al., 1999; McDonald et al., 2002). The magnitude of this redistribution depends on the intensity of tillage (Garrity, 1996) and the hedgerow spacing. The choice of hedgerow spacing depends on steepness of a slope and, in turn, determines the ratio of the area of land that is between hedgerows to that which is under hedgerows, and hence the available land for cultivation. This ratio is also important because erosion control processes, such as high infiltration rates of water, occur predominantly under the hedgerow (Kiepe, 1995), which may have significant implications for nutrient losses by leaching (see Chapter 7). The significance of the spatial variability in soil properties for experimental design was discussed by Mueller-Harvey et al. (1989) and highlights the crucial importance of including a sufficient number of independent experimental blocks, which control for variation in both site conditions and farmer practice, in order to produce conclusions of general applicability. If there is significant between-block variation, treatment effects will not be detected unless it is removed from the residual sum of squares (see McDonald et al., 2002, for an example). Location and layout of blocks should also take into account the evidence provided by Pickup

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(1985), Westoby (1987), Nortcliff et al. (1990), Busacca et al. (1993), Garrity (1996) and van Noordwijk et al. (1998) that on hillslopes there will be major transfers of nutrients within the slope between areas of net erosion (sources) and net deposition (sinks) depending on the topography of the slope (even fine-scale changes that are undetectable to the human eye). Mid-slope sites may undergo little or no net soil loss. A major impact of changes in land use, such as incorporation of trees, is likely to be on this rate of within-slope transfer. Farmers have learned to adjust their landuse practices to accommodate the effects of erosion by, for example, planting different crops in zones of net loss and net deposition (van Noordwijk et al., 1998), so that expensive technologies that minimize such transfer by surface runoff and erosion may not actually address farmers’ needs. Ramakrishnan (1990) was also concerned that, on long hillslopes, minimization of surface runoff, by increasing infiltration, may not be advantageous to overall soil fertility and agricultural productivity, because of increased nutrient losses through leaching. Therefore, the objectives of studies need to be carefully framed to distinguish net impacts of changes in land use at the individual plot scale, at the whole hillslope scale, or on the rate of discharge into rivers at the bottom of the hillslope. McDonald et al. (2002) observed that, for 14 measured soil properties, variation between blocks was much greater than that between agriculture and agroforestry treatments in an experiment in the Blue Mountains of Jamaica. This indicated that, for individual farmers operating small-scale slash-and-burn agriculture and seeking to maximize fertility of their farmland, selection of more fertile sites in which to establish new farmland was a more important decision than whether or not to adopt agroforestry practices, at least over the 5-year time course monitored by the experiment. Forest restoration through secondary succession during a fallow period may be the most effective mechanism for the recovery of soil fertility in sloping lands of the humid tropics (McDonald and Healey, 2000). However, adoption of agroforestry practices may become more important: • • •

if shortage of farmland and restrictions on tenure prevent farmers from selecting sufficiently fertile sites for new farm establishment; if long periods of cultivation lead to soil fertility falling to very low levels; or if it is the other products and services provided by agroforestry practices that are important rather than their impact on soil fertility.

Conducting experiments over a time frame sufficient to indicate long-term trends and to adequately account for the heterogeneity of the environment is essential in this type of research. Accumulation of soil material above the trees is commonly observed; hedgerows clearly do have a sieve-barrier effect that causes changes in soil

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texture over time. The higher sand concentrations above trees may partly be because sand is more easily moved down-slope than clay, which commonly leads to higher clay contents on sloping land (Fielding and Sherchan, 1999), and also reflects the fact that the hedgerow sieves trap larger sand particles more effectively than the finer silt and clay ones. The accumulation of soil above the hedgerows progressively reduces the slope angle, resulting in the formation of terraces, and, therefore, reduces runoff generation. Higher potassium and sodium concentrations under hedgerows may indicate that transport above the soil surface is a significant mechanism in their movement, either in the eroded soil that accumulates there or in runoff water, which is likely to have much higher rates of infiltration under the hedgerows (Kiepe, 1995). Ongprasert and Turkelboom (1996) indicated that on hillslopes in northern Thailand, higher soil fertility is found under contour hedgerows than between them. Such results are influenced by the relative importance of the sieve-barrier effect, which tends to concentrate fertile topsoil under the hedgerows, the transfer of prunings to the between-hedgerow zone, which tends to redistribute the fertility (Young, 1997), and the removal of nutrients from the between-hedgerow zone with crop harvests. While trees also influence soil fertility through litterfall inputs and root uptake, the spatial distribution of these two processes is highly variable. Studies of erosion within the context of agroforestry research must incorporate an additional spatial dimension. All farming systems operate within the socioeconomic and natural environment, but the forces of gravity acting upon transport processes on sloping land create complex patterns of spatial variability (Gardner, 1997). Simply measuring rates of soil movement will not be adequate for understanding the complexity of factors occurring when trees are introduced in hill environments.

Developing farmers’ criteria for evaluating soil erosion Increasingly, development-oriented research aims to be participatory, and collaborations between scientists and farmers have been particularly significant in the field of soil erosion. Studies of farmers’ knowledge about soils, erosion and fertility are still much more common than action research which seeks ways in which farmers can address soil erosion through developing their own knowledge and actions. Such academic studies are important starting points for recognizing the strengths of farmers’ knowledge, and where it differs from scientific knowledge. Results tend to emphasize the detail of farmers’ knowledge, recognizing a wide range and diversity of soils within their resource use, and the utilitarianism of their knowledge (Woodgate, 1994; Zimmerer, 1994; Carter and Murwira, 1995; Lamers and Feil, 1995). Similarly,

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farmers’ knowledge about soils and soil-related processes has been found to be quite consistent across farming communities, especially with similar agro-climatic conditions. A study in Nepal (P.K. Shrestha, unpublished) shows consistency in farmers’ knowledge about soil physical properties and their association with soil erosion and soil fertility across three different communities in the western middle hills. Some of such knowledge, however, is patchy or specific to a location, community or even individual farmers within a community due to differences in agro-climatic conditions, cultural tradition, gender and personal experience. Farmers’ knowledge of soils and soil-related processes has also been found to be complementary to scientists’ knowledge and this can help to focus further research efforts. For example, in Nepal, comparison of farmers’ perceptions of the timing of soil loss and runoff with actual measured soil loss and runoff shows that they are strongly correlated, and this may provide a shortcut to research monitoring. However, in the same areas it was found that farmers do not know about leaching losses (Shrestha et al., 2003), and it has been found elsewhere that farmers know less about below-ground processes, which are difficult for them to observe, than about above-ground processes (Sinclair and Walker, 1999). Such knowledge gaps represent problems that farmers are not likely to be able to address by themselves, and can help to identify topics suitable for participatory research. There are several reasons why it is important to involve farmers in soil erosion research, either through conducting trials on their land, inviting them to observe trials on research stations, or supporting them in conducting their own research (e.g. Okali et al., 1995). The benefits are as follows. •

•

The research will be better adapted to the environment: farmers have detailed local knowledge of what is often a highly heterogeneous environment, especially in hilly or mountainous areas where soil erosion studies are likely to be carried out. For example, studies of farmers’ knowledge in Nepal showed that agroforestry systems have quite different potentials for soil conservation in different ecological situations in the middle hills. At lower elevations, trees on cropland are acceptable to farmers, whereas at higher altitudes and on northfacing slopes, farmers find the competition between trees and crops too severe (P.K. Shrestha, unpublished). This finding can help to guide appropriate technology design. Explanations for results may be better informed by farmers’ own observations of change over many years. There are numerous documented cases where historical change known to farmers has brought about, or alleviated, the problems visible at present. Preston et al. (1997), for example, have shown that contemporary severe soil

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erosion in Bolivia is most probably due to overgrazing in the past, and that vegetation is in fact recovering at the present time. If farmers are able to observe experiments, participate in the measurements or conduct their own research, they are more likely to be convinced by the results and to interpret them in the context of their livelihood systems so that the results can be incorporated into their farming methods in appropriate ways. A tradition of participatory technology development has arisen among nongovernmental organizations and merged with farming systems research methods (Martin and Sherington, 1997), but these are particularly challenging in the context of soil conservation research, and new methods are still being developed (Lawrence, 1999; Lawrence et al., 2000). Participatory approaches can lead to shared knowledge and enhanced awareness within the community, and directly to farmers identifying indicators to measure soil erosion in their own systems, and to monitor changes in soil erosion as they or others conduct research into the effectiveness of interventions.

In order for farmers to participate in such experiments, they must use methods that are accessible and meaningful to them. Attention has focused in particular on seeking indicators of soil erosion which are identified by farmers and which can be measured by them. The relatively new group of methodologies which have grown up around participatory monitoring and evaluation (Estrella et al., 2000) are helpful here, and collective experience suggests that farmers’ indicators are likely to be visual, observable or countable but not requiring complex measurements or recording. Two case studies of indicator development summarized here contrast the priorities and approaches of farmers in systems where they see soil erosion as a low-priority problem (Andean foothills, Bolivia) and a highpriority problem (middle hills, Nepal). In both cases, farmers and scientists collaborated to monitor and evaluate the success of agroforestry and other interventions for controlling soil erosion on sloping cultivated land. In Bolivia, the communities involved are living in semiarid mountainous areas with road access to the city, resulting in migration and relatively low population densities in the rural communities. Because of this and the lower rainfall, free-range livestock are also an important component of the farming system. In contrast, in the research areas in Nepal, population densities are much higher, annual precipitation is much higher (but, as in the Bolivian case, strongly seasonal and erosive) and land use is much more intensive. As the following discussion shows, the farmers in Bolivia were initially not very interested in the soil erosion aspects of the trials, but recognized the value of this aspect with time.

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In the Bolivian case (Lawrence et al., 2000), scientists’ explanations of the value of contour hedgerows based on erosion control met with scepticism among farmers, who were, however, interested in the possibility of increasing fodder production, particularly during the dry season. As the trials developed, farmers changed the criteria that they were using to assess the trials. At an early stage, when the contour hedgerows were still small, farmers focused on potential losses brought about through the trials, and highlighted the increased labour required to prepare the land for sowing when stubble cannot be burnt; risk of losing hedgerows through browsing; and compatibility with farming practices such as ploughing by oxen. However, after one growing season, participating farmers and their neighbours began to notice both soil and moisture retention around the hedgerows, and began to express interest in establishing further trials. At this stage they were invited to explicitly identify indicators to evaluate such trials, and included soil colour, soil cover, humidity and soil content of runoff in their indicators, showing a change in perception. However, increased crop and animal production were still their priorities. In the more structured research context in the middle hills of Nepal, where farmers were specifically invited to monitor soil erosion and had more incentive to do so, given higher population densities and more intensive land use, farmers identified the following much more detailed and specific set of indicators of soil erosion (Shrestha et al., 2003): changes in the number of stones exposed on the surface; exposure of the base of terrace risers; changes in the height of the terrace risers; formation of rills or gullies; changes in soil depth; exposure of crop and tree roots; change in plant vigour and health; change in crop yield; change in outward slope of terraces; turbidity of runoff water; change in soil colour; change in soil structure. By explicitly developing soil erosion indicators with farmers, the results in Nepal provide a more detailed set of measures to analyse the success of soil conservation measures, and improve the chances of farmers and scientists communicating well. There are several lessons to be drawn from this experience. The Bolivian experience shows us that farmers’ evaluation criteria are contextspecific and may evolve during the course of a trial; they may also vary according to the season. The indicators adopted by farmers are visible and readily observable and, in both cases, farmers judge success according to their own direct experience, which often relates specifically to their own fields. The work in Nepal suggests that, although farmers look at what their neighbours are doing, unless they are actually working the soil and directly observing the colour, texture and runoff quality, they will have only a vague idea of the differences in soil erosion between plots, and between different agroforestry interventions. A trial is rarely evaluated solely on the basis of its control of erosion. Farmers are interested in looking at the effect of agroforestry (or other

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technologies) on their whole livelihood system, including labour availability, food security and resource flows for priority enterprises such as fodder for cattle. As Bunch (1999) has pointed out, rapid returns and economic benefits are more important to poor farmers with a short resource management horizon than soil erosion per se (see also Chapter 2). This last point is an important one, and not unique to the case studies described here. It is linked to a more serious point that scientists must be careful not to ignore: that, even where soil erosion appears to the outside observer to be severe, it may not be perceived as the principal problem by farmers themselves. There are plenty of case studies where soil erosion is identified as a constraint to productivity, particularly in semiarid systems, but there are others where outsiders have focused on this as a researchable constraint which can be addressed by a technological fix, whereas local residents may be aware of other priorities or sociopolitical reasons why such technological fixes will not work (Blaikie and Brookfield, 1987; Preston et al., 1997). This is why soil erosion research must take place within a social research context. The indicators that were developed with farmers in the above case studies were based on an explicit process facilitated by researchers. Farmers do not tend to consciously develop indicators, and it is sometimes the case that participatory monitoring and evaluation can place unwarranted demands on farmers, if the results are only of use to the researchers. Nevertheless, by participating in experiments which have a short-term incentive, such as fodder production, but which at the same time may reduce soil erosion, farmers may change their perceptions: those in the Bolivian case commented that they had always thought soil erosion was an unstoppable, ‘natural process’. Within 2 years of beginning contour hedgerow trials on some farmers’ fields, others were copying them spontaneously, not only for fodder production but also because soil erosion was reduced and, most importantly in these semiarid hillsides, because water was retained in the soil. It is therefore important to pay close attention to the process that is used to develop indicators. The same core process was used in the two cases described here (Table 17.1). Farmers are often unfamiliar with the concept of formal evaluation using indicators, but they are constantly observing change in the environment around them, and will be able to discuss good and bad changes easily. With careful facilitation the researchers will be able to list key words that show important change and discuss a list of them with farmers. For example, in Nepal, farmers responding to trials designed and managed by scientists noted their perceptions of good and bad aspects of the trials. These included observations about labour requirements, soil deposition and changes in crop yield, and provided a starting point for developing indicators. Once the indicators have been used in explicit evaluation and comparison of

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Table 17.1. The process of developing indicators with farmers. • • • • • •

Situation analysis, to improve understanding of the interlinkages in farming systems and to identify farmer perceptions Discussions with farmers regarding individual experiences with systems changes as a result of incorporating new farming practices Discussions with farmers (individually or collectively) regarding their expectations of incorporating new farming practices Indicator development together with farmers and researchers to show farming systems changes and impacts of new technologies Refining of systems indicators in the field by involving more farmers Use of matrix diagrams based on the identified indicators to rank and score changes

interventions or plots, their utility will be much more apparent to the participating farmers.

Research priorities Half a century of failed soil and water conservation projects in tropical developing countries demands a reconsideration of strategy (Critchley et al., 1994). Recognition of this and the changing attitude of soil and water conservation programmes since colonial times has placed much greater emphasis on people’s participation in all aspects of project design and implementation (McDonald and Brown, 2000). This has resulted in the adoption, in some areas at least, of a more holistic attitude towards the maintenance of land productivity (e.g. Shaxson et al., 1989; Douglas, 1994; Scoones et al., 1996). Much emphasis on helping farmers to improve land husbandry and less on efforts to combat erosion alone are expected to provide a more effective solution to an old problem. It has been postulated that, if farmers find it feasible and worthwhile to improve the structure, organic matter content, porosity and nutrient levels of the soil, natural fertility will be raised and soils will recuperate and runoff and erosion be diminished (Shaxson et al., 1997). It is now well recognized that the impacts of erosion are experienced differentially, and that some land managers benefit from the processes of soil erosion and runoff through redistribution of water and nutrients on their fields; for example, Stocking (1996) cites examples from Kenya and Sri Lanka, and Rakotomanana (1991) from Madagascar. Other findings suggest that rapid soil erosion may be a passing phase and that when appropriate institutional, economic and social factors are in place then higher rural population densities may be associated with investment in environmental enrichment (Tiffen et al., 1994). The dynamics of change

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Table 17.2. Considerations in defining research objectives. • • • • • •

Who is expected to use the research results? Is it necessary to convince policy makers? How heterogeneous is the area where research is to be conducted? How heterogeneous is the farming community and who makes decisions about land use/who is affected by land use? How are people (researchers and farmers) used to interacting in the culture in question? How has land use changed historically – can farmers avoid wasting time addressing causes which are historical?

and the interactions between these factors demonstrate that the assumption that more people inevitably means more erosion and greater poverty is not only simplistic but also sometimes misplaced (Blaikie, 1986; Blaikie and Brookfield, 1987). Crisis narratives have developed as a result of these simplistic assumptions leading to large-scale inappropriate and largely unsuccessful policy prescriptions, particularly in sub-Saharan Africa (Tiffen et al., 1994; Hoben, 1995; Rocheleau et al., 1995; Roe, 1995; Benjaminsen, 1998) but also in Himalayan regions (Ives and Masserli, 1989). The emphasis in monitoring erosion in agroforestry should not be restricted to monitoring soil movement but should take a holistic approach to monitoring and appraisal, combining a variety of indicators appropriate to the objectives of the research. The most valuable approach may often be to use a combination of farmers’ and scientists’ criteria, but the research must be tailored in each case to certain considerations (Table 17.2). It should be recognized that sources of error are considerable, depending on the sampling procedures, and that absolute values of soil erosion may not be meaningful. However, relative measures of soil changes, which take account of spatial patterns of relative loss and accumulation and if monitored in a participatory manner with farmers actively involved in the design and implementation of the research, can provide reasonable confidence in evaluating agroforestry innovations. Therefore, this chapter highlights some of the most appropriate methods for agroforestry research and the special methodological considerations required when trees are involved in erosion control rather than describing standard methods of measuring erosion rates that are well covered in many useful syntheses (e.g. Hornung, 1990; Lal, 1994; Hudson, 1995; Morgan, 1995; Stocking and Murnaghan, 2001).

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17.2 Quantitative Methods Runoff plots There is debate as to the value of using plot-based measurements in erosion research, as they isolate the plot from the upper hillside, and can interfere with lateral drainage patterns. However, they provide a realistic means of evaluating relative treatment effects in multivariate studies (Young, 1997; van Noordwijk et al., 1998). Historically, they are the most commonly used technique in soil erosion research as they measure runoff and soil loss directly. Runoff plots can be inexpensive and easily constructed, and thus a sufficient number can be installed to obtain a representative sampling of the major characteristics of an area such as slope, soil type, plant cover and agricultural practice. Morgan (1995) describes their construction and monitoring. Plots are rectangular with the long axis oriented up-slope, and consist of a border, an element to concentrate runoff at the bottom end, and a collector to contain the runoff and sediment produced. Plot borders may be sheet metal, wooden planks, concrete or earth bunds. The smaller the plot, the sharper the edges need to be to minimize the edge effect. Standard runoff plots are 22 m long and 1.8 m wide, but plots containing trees should be much larger, not less than 40 m ¥ 10 m (Young, 1997) to counter the problems of tree roots spreading from one treatment plot into other adjacent plots (see Section 3.2). Collectors are generally a flattened funnel with a hinged cover to facilitate cleaning. The sheet metal floor of the element is installed at least 5 cm below the surface, with a strip of sheet metal on the up-slope side flush with the soil surface to provide a sharp boundary at the plot end, which empties into collecting drums with a system of divisors to cope with large volumes of runoff. The soil loss is measured by stirring the tank, taking a sample of suspended sediment, filtering and drying, and then calculating the total loss from knowledge of the tank volume. Since most runoff plots in developing countries are simple, low-cost installations, there are many sources of error depending on the sampling procedure. Zöbisch et al. (1996) compared the manual sampling accuracy of field staff in charge of erosion plots in Kenya. The measurements displayed acceptable accuracy for measurement of runoff volumes, but the accuracy for soil loss measurements varied considerably, with an average sampling error of 41.3%. Stocking (1996) attributes this error to inadequate mixing of the solution in the tank. Even with vigorous stirring, it is difficult to get a representative sample as clays are over- and sands under-represented in the sludge samples. For large tanks, a good approach is to empty them successively, e.g. with a bucket, and collect a suspension sample at regular intervals, e.g. from each fifth or tenth bucket, depending on the total volume. After emptying the tank, the sandy sediment on the bottom of the

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tank is collected as a separate sample. After drying all samples, the quantity of eroded soil is calculated by multiplying the average concentration of suspended soil per unit volume of water from the suspension samples with the total water volume in the tank and adding to this the bottom sediment (T. Morshäuser, unpublished). Runoff plots have been used in several agroforestry studies (e.g. Hurni and Nuntapong, 1983; Maass et al., 1988; Lal, 1989; Agustin and Nortcliff, 1994; Alegre and Rao, 1996) and modifications have included use of Gerlach troughs (Maass et al., 1988; McDonald et al., 2002) or catchpits (Hellin and Larrea, 1997). These are described below. Difficulties arise in scaling up results from such plots to farmers’ fields, hillsides or catchments, so, for a better understanding of erosion as a multiscale process, other indicators of processes within standard plots should also be observed (see subsequent sections). This is particularly important as agroforestry research evolves from a focus on field-scale technologies to a more complete consideration of landscape-scale processes and constraints involving a farmer-led process of technology development (van Noordwijk et al., 1998).

Back-sloping terraces: a special case Runoff plots were developed for uniform slopes with gentle gradients in the USA (Wischmeier and Smith, 1978). Hillside terraces are far smaller than fields, and a standard runoff plot will therefore encompass upwards of three terraces. As yet, there is no standard means of quantifying runoff and erosion losses from such terrain (Hudson, 1992). With the backsloping terraces common throughout South-east Asia, bounded plots will dissect lateral drainage paths (toe-drains), and the collecting drums may effectively collect only from the lowest terrace, giving artificially low values (Bruijnzeel and Critchley, 1996). Bruijnzeel and Critchley (1996) have suggested the concept of a natural boundary erosion plot based on a singleterrace unit (bed and up-slope riser) with its actual boundaries (Fig. 17.1). The toe-drain is located at the foot of the riser, so runoff and eroded material are channelled out of the individual terrace unit and can be conveniently monitored at the outlet. This will also collect uncontrolled surface run-on or subsurface water input from upper slopes (Gardner et al., 2000), so such measurement devices should be installed on several terraces in different positions within the slope to obtain an overall picture. Bruijnzeel and Critchley (1996) suggest another, specific metholodogy to determine the source of the eroded material. This involves installing a set of removable terrace riser troughs, which collect runoff, sediment washed down from the riser face and, presumably, any subsurface water flow emerging from the riser face (Fig. 17.1).

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Fig. 17.1. Natural boundary erosion plot with one terrace riser trough (TRT) in place (reproduced with permission from Bruijnzeel and Critchley, 1996).

This area of research will become more relevant to agroforestry research if the objective is to control riser erosion and increase the productivity of terraced systems, e.g. by inclusion of perennial fodder grasses and legumes, with stabilizing fodder trees planted on the top of the riser (Shrestha et al., 2003).

Gerlach troughs This method is described in detail by Morgan (1995). Gerlach troughs are simple metal gutters, usually 0.5 m long and 0.1 m broad, closed at the sides and fitted with a removable lid. An outlet pipe runs from the base of the gutter into a covered collecting vessel (Fig. 17.2). They are normally sited at different slope lengths and the collecting area assumed to be equal to the width of the trough times the slope length (Fig. 17.2). On nonuniform slopes, the gutters should be sited to collect down-slope and not lateral drainage. The lack of plot boundaries eliminates any edge effects. These gutters can therefore be used without runoff plots (e.g. McGregor, 1988) to investigate the interaction of soil status and soil erosion with land use. They provide a relatively efficient means of covering a range of conditions within a site. They have been used to collect runoff and eroded soil from small bounded subplots nested within a treatment plot (e.g. Ross et al., 1990) but, clearly, this is not an applicable approach in plots containing trees. Small plots will allow investigations into infiltration and the effects of rain splash but are too short for studies of overland flow except as a transporting agent for splashed particles (Morgan, 1995).

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Fig. 17.2. Gerlach troughs (reproduced with permission from Morgan, 1995).

Catchpits Catchpits are collecting pits that are dug into the ground at the bottom of the study area and lined with plastic or concrete to render them impermeable and that fill with runoff water and eroded soil. Catchpits can be effective in measuring amounts of sediment lost and runoff volumes if

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the reservoir is of sufficient capacity, although smaller pits that only catch an unknown proportion of the sediment can still be used to obtain comparative information. Pits of around 10 m3 have been used in Thailand (Hudson, 1995). Obviously, durable plastic is required that will not be holed by soil fauna or degraded by sunlight. Hellin and Larrea (1997) used catchpits lined with plastic that had been punctured with small holes so that runoff water slowly dissipated and the sediment load was retained until end-of-season measurement. They found soil loss to be within the same range as that recorded by barrels below similar plots. Catchpits have the added advantage of providing a visual display for farmers of the effectiveness, or not, of the practice under test.

Stakes and pins Repeated measurement of the height of the ground surface at stakes or erosion pins has been used to estimate erosion. Instrumentation consists of a long nail or stake and a large washer. Stakes should be made from metal or another resistant material as wooden stakes are rapidly attacked by termites under tropical conditions. At the time of installation, the nail is driven into the soil, with the washer lying on the soil surface. The distance from the head of the nail to the top of the washer is then measured (Fig. 17.3a). Erosion moves material from around and beneath the washer. Remeasurement of the distance between the top of the nail and the top of the washer provides a measure of the rate of erosion in the intervening period (Fig. 17.3b). If the washer has protected the soil from raindrop impact, so that it now stands on a pedestal, the pedestal must be removed before measurement, so that the washer lies at the general ground surface. The advantage of using the washer is that it gives a firm surface from which to measure. The washer can also be removed between two measurements. Marking the pins with bright paint aids relocation. Such measures are cheap and easy to install over a wide range of conditions. A disadvantage of the method is that it cannot be used on tilled soil because of the change of soil surface caused by tillage and the subsequent resettling of the soil. Instead of measuring soil removal by erosion, pins can also be used to measure soil deposition. In a sloping area in central Togo, G. Schroth (unpublished) used the method to determine patterns of sediment deposition in strips of trees and hedgerows which had been planted on the contours to collect soil lost from adjacent tilled crop fields where the pin method could not be used. Pins could be a cost-efficient method to determine patterns of net erosion and deposition over entire slopes (see Section 17.1). Instead of measuring directly around the pins, a cable or other rigid device can be placed between two pins or stakes. With this improvement

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Fig. 17.3. Measurement of erosion and deposition at stakes: (a) installation; (b) remeasurement (reproduced with permission from Dunne, 1977).

of the pin method, Sims et al. (1999) evaluated the effects of barriers on soil retention with the help of permanent benchmarks above and below them. Measurements were made to the soil surface from a taut cable tied to the benchmarks, which indicated the accumulation or loss of soil before and after each rainy season. This was combined with annual measurements of slope of the interbarrier soil to indicate changes in microtopography.

17.3 Qualitative Methods Tree root exposure or soil pedestals It is sometimes possible to reconstruct the recent erosion history of an area from truncations of soil profiles, from the height of residual soil pedestals, or from the exposure of tree roots. Normal spatial variations of soil profile depth with slope gradient must be taken into consideration. If soil erosion is rapid following the destruction of vegetative cover, remnants of the former surface may be left. A ruler, tape or frame laid across the former surface can be used as a reference from which to measure the average depth of erosion (Fig. 17.4a). The exposure of tree roots can also be measured (Fig. 17.4b). However, some trees, even on undisturbed sites, grow with part of their roots above the ground surface. So, before such measurements are made, trees of the same species in neighbouring areas should be investigated.

Changes in soil microtopography Evidence of the breakdown of the surface soil structure and its subsequent transport across the soil surface can be seen in the change in microtopo-

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Fig. 17.4. (a) Measurement of erosion between vegetated remnants of former soil surface; (b) measurement of erosion around tree roots (reproduced with permission from Dunne, 1977).

graphical features of the soil surface. These include the breakdown of clods formed by ploughing; the filling of ploughed depressions by soil loosened from the clods and from the rest of the soil surface; and indicators that water has flowed across the soil, such as flowlines and rills. Ranking schemes can be developed to undertake series of structured field observations of the evidence for soil loss provided by these features (Bergsma, 1992). Stocking and Murnaghan (2001) suggest developing a context-specific ranking scheme which, if used consistently, can be a good way of combining indicators to give a more comprehensive view of land degradation. For example, to assess rill erosion, a scale of 0–3 indicates the development from no rills present to the presence of deep rills (up to 300 mm depth) and/or rills affecting more than 25% of the surface area. The FAO (1976) advocates wider ranking for the same parameter with a scale from 0 to 14, which describes a range from no visual evidence of rills, to rills present 7.5–15 cm deep at intervals less than 1.5 m. Utilization of numerically wider ranges has the added advantage of being more sensitive to the determination of statistical significance. The FAO (1976) and Bergsma (1992) apply such rankings to a range of topographical features including flow patterns, gullies, surface litter accumulation and depressions. Sources of error can include classification of the features, a low frequency of observations, and mislocating the same location for repeated recordings (Bergsma, 1992). Special attention should be given

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to signs of runoff and transport of eroded soil at the field boundaries, where soil is effectively lost from the field, i.e. on the down-slope side in all fields and on the lateral sides in contour-ridged and terraced fields.

Stable isotopes Caesium-137 is a by-product of the nuclear bomb testing that occurred in the 1950s and 1960s. It has a relatively long half-life of 30.17 years and does not occur naturally in the environment, so can be used as an indicator of disturbance if comparison is made between undisturbed areas and those subject to perturbation by erosion. The technique has been applied successfully to terraced areas in China and Nepal (Quine et al., 1992; Zhang et al., 1994; Gardner et al., 2000). Gardner et al. (2000) noted very high spatial variability, with the worst cases indicating losses roughly equivalent to 40–50% of the top 10 cm of soil over the 40-year period. They concluded that there was no systematic spatial pattern of net loss or accumulation down-slope associated with natural convexities or concavities in the slope topography, and that the differences were more likely to reflect localized pathways of preferential drainage and water movement. Their results highlight the processes of redistribution of soil on a hillslope scale, and indicate the usefulness of this technique as a relative tool for qualitative assessment of soil erosion to indicate patterns of spatial loss and accumulation. The extension of the technique to provide quantitative estimates of loss remains to be developed (Walling and Quine, 1990). Estimates of soil loss using caesium-137 should be compared with other means since they may be subject to errors arising from uneven mixing of the cultivated layer and preferential adsorption of the isotope on to clay particles during the processes of erosion, transport and deposition (Morgan, 1995).

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Index

Acacia spp., 261 Acacia auriculiformis, 194 Acacia mangium, 138 Acacia nilotica, 194 Acacia saligna, 154, 246, 256 and reduced leaching, 156 Acacia senegal, 168 Acacia tortilis, 168 Acer saccharum, 234 acid soils, 127–130 and soil organic matter, 78 actinomycetes, 259 actinorhizal symbioses, 259 Africa, 2 savanna soils, 83 soil rhizobial inoculation, 264 Africa, East, 114 Africa, sub-Saharan, 6 erosion control, 335 rainfall and nutrient gain, 181 Africa, West alley cropping trials, 110 atmospheric nitrogen, 181 and fire damage, 184 mineral leaching, 153 nutrient deposition, 182 nutrient pumping, 168

and reciprocal relations, 15 ridging, 58 savanna soils and carbon loss, 80 savannas, 2 soil degradation, 192 aggregate stability measurement, 196–198 and soil organic matter, 204–207 agricultural markets agroforestry product sales, 24–25 as decision-making factor, 15 agroforestry classification of practices, 4, 5, 6 definition, 3, 5 and improved crop yields, 23–24 and landscape design, 5–6, 45 legume trees and nitrogen efficiency, 95 as natural capital, 32–37 and phosphorus methods, 112–121 practice matching to sites, 103 product sales, 24–25 range of benefits, 2–3 and rhizosphere processes, 290–291 and risk management, 27 trial hierarchy of levels, 47–48 423

424

agroforestry processes and economics, 36–37 and nutrient cycles, 95–98 agroforestry trials break-even point, 29–30 long term and replicates, 47 Albizia guachapele, 109 Albizia niopoides, 109 Alfisol, 114, 118, 122, 191, 192, 279 alley cropping, 27, 28, 31–32 in moist savanna, 110 allophane, 169 aluminium, 112, 113 measurement, in soil, 129–130 oxides, 78 and sulphate leaching, 152–153 toxicity, 93, 262 toxicity diagnosis, 128–129 Amazonia, 199, 201, 246, 310 abandoned pastures, 127 earthworms, 315 leaching and tree spacing, 157 nitrate sorption, 169 nutrient loss by fire, 184 Oxisol and soil organic phosphorus, 117 soil acidity, 130 soil degradation and mechanical clearance, 192 and sulphur availability, 123 sulphur leaching, 153 and time domain reflectometry, 218, 219 tree root measurement, 254 water efficiency, 212 America, Central, 264 agroforestry practices NPV, 27 nutrients from tree litter, 132 America, South, 264 ammonia, 182 ammonium, 104 animal residues, 83 antibiotics, 275 ants, 161, 306–307, 309 structures, 310, 322 apatite, 113 Arachis hypogaea, 278 arbuscular mycorrhizal fungi

Index

and improved infection resistance, 275 isozyme analysis, 284–285 molecular identification, 285–286 non-vital staining, 281–282 and nutrient transfer, 275–276 and phosphatase activity, 273, 277 and plant water uptake, 274 population management, 278–279 quantification and identification, 281 quantification using spores and sporocarps, 286 and rhizosphere processes, 290–291 and rhizosphere zones, 295 soil as an inoculum, 280 species variations, 277–278 visual evaluation methods, 282 see also mycorrhizal fungi Asia, South-east grassland fires, 1 imperata grasslands, 127 Ateleia herbert-smithii, 109 atmospheric nutrient deposition, 95, 104 atmospheric nutrient exchange, 181–189 Australia ferralitic soils and aggregate stability, 207 fodder trees and water use, 217 nitrate accumulations under crops, 173 soil analysis, 89 soil degradation, 192 Azadirachta indica, 168, 234

Bactris gasipaes, 157, 175, 212, 313, 315 Belize, 169 Benin, 155 Bertholletia excelsa, 157, 313, 315 Betula spp., 148

Index

BIOASSESS programme, 321 biodiversity, 5–6 conservation and agroforestry, 315–316 as monetary benefit, 25, 26 biomass, 77 decomposition and nutrient supply, 131–150 managing litter and prunings, 140–143 measures of resource quality, 148–150 nutrient cycling, 132–134 nutrient leaching, 134–135, 154 nutrient release prediction, 135–138 optimum use decision trees, 137 tree production factors, 133 and tree pruning, 102 biomass decomposition process, 136 management, 138–139 methods, 143–148 Bixa orellana, 157, 313, 315 Bolivian erosion control, 330–333 book objectives and structure, 9–11 boron, 128 leaching, 153 boundary plantings, 20, 102, 224 Brachystegia spp., 269 Brazil adsorption measurement by radioisotope, 119 nitrogen mineralization measurement, 107 nutrient pumping, 169 Oxisol macroaggregation, 206 Oxisols and fertilizers, 93 phosphorus fertility decline, 118 soil analysis, 89 soil nitrogen chemical fractionation, 111 bromide, 163, 164 Burkina Faso parkland water use, 233 soil organic matter analysis, 91 tree root growth, 173 Burundi, 154, 155

425

Caesalpinia coriaria, 109 Caesalpinia eriostachys, 109 Caesalpinia velutina, 109 Caesalpinioideae, 260 Cajanus cajan, 44, 134, 138, 265 calcium, 113, 114, 128, 129, 163, 178 leaching, 153, 169 methods, 125–127 California, 177 Calliandra calothyrsus, 23, 140, 170, 264 Camellia sinensis, 128–129 Cameroon, 269 carbon, 60, 121 loss, 80 from roots, 300 management index, 84, 85 mineralization, 311 organic, 78, 81–82, 83, 89 methods, 86 sequestration, 25 soluble, 138, 149 proximate fractions analysis, 150 transfer, 275–276 carbon dioxide, atmospheric, 79 Caribbean agroforestry practices NPV, 27, 28 Cassieae, 260 Casuarina spp., 259 Casuarina equisetifolia, 275 cereal straw, 80, 138, 140 and sulphur, 122–123 Chamaecrista, 260 charcoal, 81 Chinese agroforestry practices NPV, 27–28 chitin assay of mycorrhizal fungi, 283 chloride, 163, 164 chlorophyll synthesis, 121 chromatographic analysis, 88 Chromolaena odorata, 109 Citrus spp., 177 Citrus aurantifolia, 279 clean-weeding, 238 Cocos nucifera, 157, 177 Coffea arabica, 177

426

Colombia, 177, 306, 309, 311 Oxisol and soil organic phosphorus, 118 compost, 79–80 and sulphur, 122 conifer plantations and phosphorus, 120 contour hedgerows, 20, 27, 96, 102, 332, 333 and erosion control, 327, 328–329 Cordariocalyx gyroides, 269 Cordia alliodora, 126, 155, 216 Cordyla pinnata, 277 cost–benefit analysis environmental, 34, 35–36 environmental and social benefits, 23–26 landscape-scale natural capital, 32–37 monetary costs, 20–21 opportunity costs, 21–23 private, 19–31 result interpretation, 29–31 sensitivity analysis, 28 time horizon, 29–30 social, 34–35, 36 time horizons, 26–28, 29 Costa Rica, 6, 177, 216, 254 farmers’ risk management, 27 leaching from monocrops, 155 shade trees, 236 Côte d’Ivoire, 310, 311 earthworm communities, 314 nitrogen mineralization measurement, 108–109 nutrient deposition, 183 soil regeneration, 192, 194 soil water study, 233 crop prices, 28 crop residues, 77 crop rotation and fallows, 102 crop yields and agroforestry, 23–24 crops and nutrient-use efficiency, 94–95 crown leaching and nutrient release, 168, 169 crown pruning in dry weather, 214 Cupressus lusitanica, 123 Cussonia barteri, 233

Index

Dactyladenia barteri, 174 data analysis, 39–76 exploratory analysis, 62 multiple outputs, 62 objectives of trial, 61 preparation, 61–62 repeated measurement analysis, 63–64 replicate samples, 242 spatial structure, 67–76 spatial variations, 64–65 timescale of experiments, 62–63 variance and regression, 63 decision-making processes ecological parameters, 16–18, 19 economic parameters, 16–18, 19 farmers’ participation in research, 329–334, 335 and social norms, 15, 17, 18 socio-political parameters, 16–18, 19 deforestation, mechanical, 192, 205 Desmodium intortum, 154, 155 Desmodium ovalifolium, 205 Desmodium rensonii, 269 dew, 182 Dialium guineensis, 277 Dicorynia guianensis, 315

earthworms casts, 310, 311, 321–322 and phosphorus, 120 functional categories, 308, 309 and organic matter assimilation, 314–315, 317 and soil compaction, 316 and soil mineralization, 311 sphere of influence, 305, 306–307, 308, 309 ecological economics, 14 economics and soil fertility management, 13–37 ecosystem engineers, 305, 310–311, 316–317 biostructures, 306–307, 311 ecosystem services, 32–37 ectomycorrhizal fungi, 271

Index

and increased resistance to infection, 275 isozyme analysis, 284–285 and mineral acquisition, 272 and nitrogen/phosphorus poor soils, 277 nutrient transfer, 275–2776 and organic acid exudation, 273 phosphatase activity, 273–274 and plant water relations, 274 population inoculation and management, 278, 280 quantification by PCR, 283 quantification using spores and sporocarps, 286 and rhizosphere processes, 290–291 sample collection and storage, 281 species variations, 277–278 visual evaluation, 282, 283 see also mycorrhizal fungi Ecuador, 6 Elaeis guineensis, 58, 170, 177, 192 and endophytic nitrogen-fixing bacteria, 261 root analysis, 243, 244 environmental impact on decisionmaking processes, 16 erosion, 79, 80, 96, 120, 128, 325–343 in agroforestry, measuring, 327–329 control, 6, 25 farmers’ criteria for evaluation, 329–334, 335 with trees and shrubs, 326–327 research back-sloping terraces, 337–338 catchpits, 339–340 Gerlach troughs, 38–339 priorities, 334–335 qualitative methods, 341–343 quantitative methods, 336–341 runoff plots, 336–337 soil microtopography changes, 341–343 stable isotopes, 343 stakes and pins, 340–341 tree root exposure, 341

427

Erythrina poeppigiana, 126, 216, 262, 264 Ethiopia, 123 Eucalyptus spp., 138, 290 Eucalyptus deglupta, 236 Eucalyptus globulus, 254 Eucalyptus grandis, 272 Eucalyptus maculata, 272 European forests, 107 experiment design, 39–76 control treatments, 42 controlling variables, 43 factorial treatment structure, 42 fallows, 48–52 measurements, 52–53 objectives, 40–41 quantitative levels, 42–43 software, 65–66 time and spatial structure, 43–45 treatment definition, 41–43 layout, 43–45 controlling variables, 46–47 insurance and long-term trials, 47 random blocks, 46 replicate numbers, 46–47 experimentation costs on farm, 20 on station, 21

Faidherbia albida, 21, 168, 213–214, 233, 269 farm subsidies, 30 farmers, resource-poor, 3 farmers’ criteria for erosion control, 329–334, 335 farmers’ decision-making processes, 14–18 contextual hierarachy, 17, 18 at farm scale, 18–32 scientists’ reductionism, 16–17 farmers’ views, 8–9 ferralitic soils, 192, 199, 207 Ferralsols, 78, 84, 87

428

fertilizers, 14, 153–154, 169, 247 costs, 20 and phosphorus availability, 112, 114 and root growth, 238 and soil organic matter, 80 fire, 1, 2, 94, 96, 104, 117 and nutrient loss, 183–184, 188–189 Flemingia macrophylla, 219, 269 fodder, 2, 6, 132, 140, 170 and deposition of excreta, 168 and erosion control, 332, 333 fog and cloud belts, 182 food production sustainability, 32–33 forest gardens, 59 French Guiana, 315 fuelwood, 170, 238 fulvic acid, 89

gas chromatography and sulphur, 122 geostatistics, 241 alternative methods, 75–76 analysis procedure, 70–72 kriging, 72–75 nugget variances, 70, 71–72 principles, 68–70 regionalized variables, 69–70 sampling grids, 71 semivariograms, 70–72 simulation, 75 software, 76 uses, 67–68 Ghana, 177 Gliricidia spp., 279 Gliricidia sepium, 109, 110, 138, 177, 264, 265 global scale agroforestry benefits, 32–37 Glomus aggregatum, 277 Glomus deserticola, 274, 275 Glomus fasciculatum, 275 Glomus intraradices, 277 Gmelina arborea, 126 Gossypium hirsutum, 177 government policy and farmers’ decision making, 15, 17

Index

green manure, 261 and aluminium toxicity, 130 green revolution, 7 Grevillea robusta, 241

Hawaii, 254 heavy metal pollution, 261 hedgerow intercropping, 20, 25, 28, 102, 126, 132, 144, 156, 217, 219 experiments, 56 and improved soil structure, 192 and tree roots, 174, 179 Hedley fractionation method, 117, 118 Heuca brasiliensis, 276 hill slopes, 6 and erosion, 327–329, 336–338 HIV/AIDS and labour availability, 21 homegardens, 27 human population growth, 2, 210 humic acids, 89

India, 177 farmers’ risk management, 27 forest soil transportation, 16 hedgerow intercropping, 217 soil improvements, 194 sulphate sorption in acid subsoils, 122 infiltrometers for boreholes and wells, 229 double-ring, 227–229 sprinklers, 229 tension or disc permeameters, 230–231 invertebrates classification systems, 317 and litter decomposition, 147–148 ion chromatography and sulphur, 122 iron, 112 oxides, 78 toxicity, 128

Index

isotopes, 266–268 carbon, 90–91, 145 hydrogen, 164 radioactive, 119, 124–125, 127, 163–164, 176–177, 179 stable, 163–164, 178, 179, 214–215, 232–234, 343

Jamaica, 328 Java, 236 Julbernardia spp., 269

kaolinite, 78, 169 Kenya, 177, 224, 334 alley cropping, 28 fallows and nitrate use, 172 hedgerow intercropping, 217 improved fallows time horizon, 29 intercropping tree species, 43 labour sexual restrictions, 22 land use tradition, 22 eaching management, 154 nitrate sorption, 169, 171 plant water use measurement, 219 soil water profile measurement, 216 trees and soil evaporation, 213 western, 25 Kjeldahl procedure, 105, 269

labour costs, 14, 15, 20, 21–22, 28 land degradation, 127–128 disputes and trees, 19 resources, 15 land-use change and soil organic matter, 79–80, 91 landscape patches and agroforestry interfaces, 45 Latin American agroforestry systems NPV, 28 leaf mulch

429

and phosphorus, 114 and sulphur, 122 leaves and nitrogen concentrations, 133 Leguminosae, 2, 8, 102, 132, 259–261 affected by management, 262 nitrogen fixation determination methods, 265–269 nodulation and nitrogen fixation, 260–261 and nutrient cycling, 95 response to stress, 261–262 rhizobial inoculation requirements, 264 and rhizobial nodulation, 263–264 Leucaena spp., 279 Leucaena collinsii, 109 Leucaena diversifolia, 262 Leucaena leucocephala, 21, 23, 109, 148, 154, 155, 219, 264, 265, 273 lignin, 85, 102, 136, 137 lime, 127, 130 and potassium uptake, 125–126 lithium, 127, 163, 179 litter, 55, 77, 84, 213 and biomass input methods, 140–143 decomposition, 276–277 bags for collecting, 143, 145, 147 baskets for collecting, 144 constants, 148 environmental factors, 148 leachate monitoring, 146 and leaf waxiness, 149 and microorganisms, 147–148 and nutrient release, 168, 169 quantification, 142–143 selection and treatment, 146 standards, 148 traps, 141–142 unconfined, 144, 147 and improved soil structure, 192, 194 as invertebrate habitat, 313–314 and mobile nutrients, 133

430

litter continued and nutrient release, 138 production and quality, 101–102 and soil acidity, 139 and soil organisms, 139–140 litterbags and invertebrates, 322–323 Lixisol, 108–109, 110 lysimeters and phosphorus measurement, 120

machinery resources, 14, 15 MACROFAUNA database, 320 macropores, 151–152, 156, 165, 205 and improved soil structure, 193–194 and water infiltration, 230 Madagascar, 334 magnesium in soil, 128, 129, 178, 272–274 leaching, 153, 169 methods, 125–127 and tree biomass, 133 Malawi, 269 intercropping, 28 maize intercropping costing, 21 Malaysia, 177 Malus domestica, 177 manganese, 128 leaching, 153 Mangifera indica, 177, 269 Manihot esculenta, 278, 279 manure, 79–80, 93–94, 115, 140, 154 and sulphur, 122, 123 Medicago sativa, 178 Mehlich procedures, 113, 116 microbial biomass, 80, 83, 298 estimation, 89 and nitrogen, 111–112 and phosphorus adsorption, 116 microbial soil changes, 60 microorganisms, 79 and litter, 139–140 and litter decomposition, 147–148 and soil solution measurement, 160–161

Index

microorganisms’ functional domains, 304–309 Mimosoideae, 260–261 mineralization flushes, 90, 107, 152–153, 153 minirhizotron observations, 146 modelling bioeconomic processes, 31 economic processes, 31–32 soil organic matter dynamics, 81–83 molybdenum, 128, 261 Mozambique, 19 mulch, 213 and aluminium toxicity, 130 and improved soil structure, 192 Musa spp., 177 mycorrhizal fungi, 248, 271–287 chitin assay, 283 immunochemical detection methods, 284 and improved soil structure, 193 isozyme analysis, 284–285 laser scanning confocal microscopy, 283 and mineral nutrient uptake, 286–287 molecular identification, 285–286 nitrogen transfer from nitrogenfixing plants, 287 and nitrogen uptake, 104 and phosphorus availability, 120 quantification using spores and sporocarps, 286 research needs, 279–280 see also arbuscular mycorrhizal fungi; ectomycorrhizal fungi

natural capital, 32–37 nematodes, 275 Nepal, 335 erosion control, 330–333 net present value (NPV), 20 break-even point, 29–30 and time horizons, 26–28 New Zealand, 219 Niger, 182

Index

Nigeria, 224, 279, 314 fuelwood production, 238 improved soil structure, 193 Lixisol and nitrogen uptake, 110 plant water use measurement, 219 savanna soils and organic phosphorus, 117 soil degradation, 191 soil organic matter formation, 85 nitrate, 104 leaching, 69, 99, 152, 171–173 sorption, 169 nitric acid, 182 nitrite, 104 nitrogen, 60, 78, 85 atmospheric deposition, 181–182 availability and agroforestry, 103 availability from biomass, 135, 138 chemical fractionation, 111 deficiency, 104 dynamics, 82 fertilizer, 80 leaching, 171–173 methods, 104–112 and microbial biomass, 111–112 mineralization, 105–106, 106–109, 107–112 organic, 104 fractionation, 109–110 labile, 109–110 physical fractionation methods, 110–111 resource quality measurements, 149 total, 105 and tree biomass, 133 uptake, 272–274 nitrogen fixation, 6, 101–102, 104 biological, 259–270 environmental limitations, 261–262 from green manure, 261 and nodulation in Leguminosae, 260–261 management effects, 262–264

431

testing in controlled conditions, 267 by empirical observation, 265–266 estimating totals, 270 by excavation of nodules, 265 in field settings, 268–269 isotope-based methods, 266–268 by transport products, 269–270 nitrogen-fixing species root analysis, 246 nitrogen-fixing trees, 133 and nitrogen availability, 262 nodulation affected by management, 262 in Leguminosae, 260–261 response to stress, 261–262 rhizobia, 263–264 nodules, isolating rhizobia from, 265 nuclear magnetic resonance spectroscopy, 88, 120 nutrient atmospheric deposition methods artificial filters, 186 canopy balance method, 186–187 micrometeorological methods, 185 plant surface washing, 185–186 sample collection and analysis, 188 nutrient availability, 93–130, 245 from biomass, 131–150 sampling frequency, 60–61 and soil organic matter, 78 nutrient capture, 167–179, 236 from subsoil, 168–173 nutrient compartments, 97 nutrient cycles and agroforestry, 95–98 in biomass, 132–134 optimization, 101–103 and rhizosphere processes, 290 in space and time, 102 and tree fallows, 103 and tree pruning, 102–103 nutrient deposition atmospheric, 181–183 from dew, 182

432

nutrient deposition continued from dust and aerosols, 182–183 from fog, 182 nutrient efficiency, 94–95 and tree–crop interactions, 98–101 nutrient horizontal redistribution, 173–174 nutrient input methods from biomass, 140–143 nutrient leaching, 78, 103, 104–105, 127, 129, 134–135, 151–166, 171–173, 326 and climatic factors, 151–152 and groundwater pollution, 106–107 management for reduction, 153–154 from organic sources, 154–155 and soil conditions, 151–152 substance susceptibility, 152–153 sulphur, 122 traced by dyes, 165–166 tree effect, 126, 155–158 and tree spacing, 157 nutrient leaching methods lysimeters, 158–159, 161–162 resin cores, 158–159, 162–163 suction cups, 158–161 tracers, 163–166 nutrient loss through fire, 183–184, 188–189 nutrient pools, 96–97, 131 nutrient pumping, 168–173 nutrient release management, 138–139 monitoring biomass leachate, 146–147 predicting, 135–138 synchrony and synlocation management, 134–135 nutrient sequestration and tree pruning, 102 nutrient transfers, 98 and mycorrhizas, 275–277 nutrient uptake and root moisture exudation, 291 sampling, 56

Index

nutrient uptake measurement lateral distribution, 179 by radioisotope, 176–177 by rare elements, 178–179 stable isotopes, 178 tracer methods, 174–175 vertical distribution, 175–176 nutrition benefits from mycorrhizal fungi, 272–274

Olsen procedure, 113, 116 Oxisols, 122, 125, 127, 154, 155, 157, 169, 192, 201, 206, 218, 221, 310 and fertilizers, 93 and soil organic phosphorus, 117, 118, 119

Panama, 233 Papilionoideae, 260–261 Paraserianthes falcataria, 236 Parasponia, 260 Parkia biglobosa, 173, 277 parkland systems, 2, 102, 132 Passiflora edulis, 315 Peltophorum dasyrrhachis, 138, 156 Pennisetum glaucum, 279 perennial crops with trees, 27 phenols, soluble, 149 Philippines, 177, 194 farmers’ risk management, 27 phosphatase, 273 phosphate, 113 resistance to leaching, 153 phosphorus in soil, 60, 78, 94, 128, 272–274 adsorption, 114–115, 116, 119–121 atmospheric deposition, 181–182 deficiency symptoms, 113 dynamics, 82 extraction procedures, 113–114 leaching, 153 methods, 112–121 and microbial biomass, 116, 118–119

Index

and mineralization, 118 and physical fractionation, 120 pools, 116, 117, 118, 119, 290 resource quality measurements, 149 sequential fractionation methods, 115, 116, 117–118, 120 spatial pattern sampling, 58 and tree biomass, 133–134 Picea abies, 272 Pinus elliottii, 179 Pinus kesiya, 194–195 Pinus radiata, 257 Pinus rigida, 273 Pinus sylvestris, 272, 276 and mycorrhizal interaction, 276 plant–pathogen interactions, 275 plant residues, 80, 83, 104 plant root sampling for mycorrhizal fungi, 281 polyphenols, 85, 102, 136, 137 polysaccharides and soil structure stabilization, 206 Portugal, 254 potassium in soil, 128, 129, 153, 163, 179 atmospheric deposition, 181–182 methods, 125–127 and sulphate leaching, 152–153 and tree biomass, 133 Prosopis juliflora, 194 Psidium gujava, 177 Pueraria phaseoloides, 205, 276, 279, 313, 315 pyrolysis, analytical, 88

Qualea spp., 315

radiation, 213 rainfall and atmospheric deposition, 182 input and use, 209 and nutrient release, 134–135 rape leaves and sulphur, 123 rhizobia, 260, 261–262, 263 isolating from nodules, 265

433

microbiological study methods, 265 requirements for inoculation, 264 rhizoplane, 296 rhizosphere processes, 289–301, 305 boundary definition, 292 ecology of non-crop plants, 293 importance in agroforestry, 290–291 mathematical modelling, 300–301 methods for biological activity, 298–299 and microbial interaction, 294 and phosphorus, 120 root carbon loss, 300 soil chemistry methods, 297–298 soil sampling, 294–296 spatial and temporal measurement, 292–293, 299 study in laboratory mesocosms, 293–294, 296 study problems, 292–294 and sulphur mineralization, 122 zones, 295 ridging, 58 risk management, 25, 27 root hydraulic conductivity, 274 root litter, 77 rubidium, 127, 163, 179 Rwanda, 28

Saccharum officinarum, 261 Sahara, nutrient deposition, 182, 183 Sahel, 234 salinity, 261 savanna soils, 78, 83, 85 and agroforestry for fertility, 103 dispersion techniques, 88 mineral leaching, 153 and organic phosphorus, 117 sediment transport rates, 325 seed costs, 20 Senegal nutrient pumping, 168 Senna spp., 279 Senna siamea, 173, 260 and calcium accumulation, 126 Senna spectabilis, 41, 260

434

Sesbania rostrata, 263 Sesbania sesban, 44, 110, 120, 170, 264 shade as on-farm benefit, 26 and perennial crops, 20 shade trees, 79 in coffee/cocoa plantations, 2, 102, 103, 126, 132, 236, 249, 254 shoot pruning, 102–103, 247, 248, 249 and nitrogen fixation, 262 and root growth, 238 sodium measurement, 187 and sulphate leaching, 152–153 soil acidity, 93 and litter, 139 measurement, 127–130 soil aggregation, 196–198, 204–207, 310 soil bulk density methods alternative methods, 196 clod method, 196 cores, 195–196 soil chemistry and rhizosphere processes, 297–298 soil cores, 108, 195–196, 240–242, 244, 248–249 soil degradation, 7, 128 and earthworms, 316 soil dispersion, 87–88 soil enzymes, 90 soil fertility and agroforestry choices, 328–329 decline following fallows, 50–52 and destruction of soil fauna, 303 improvement, 25, 26 as monetary benefit, 25–26 and organic matter, 78–79 recovery during fallows, 49 spatial pattern sampling, 57–59 and trees, 6–9 soil fertility management decision-making processes, 14–18 economic aspects, 13–37 economic processes, 36–37 short-term benefits, 30

Index

soil hydraulic properties methods, 225–226 infiltrability and saturation conductivity, 226–229 and pedotransfer functions, 231 unsaturated conductivity, 230–232 soil improvements as monetary benefit, 25 soil macrofauna, 303–323, 304 and agroforestry practices, 312–318 biogenic structure collection, 321–322 excrement, 305, 321–322 handsorting, 319 identification and data storage, 319–320 and improved soil structure, 194 pitfall traps, 321 sampling designs, 318 and water conservation, 213 see also soil macroinvertebrates soil macroinvertebrates, 303–323 effect on soil processes, 309–311 and litter, 313–314 manipulative experiments, 322–323 research needs, 316–318 and tree species, 315 see also soil macrofauna soil management and organic matter, 83–84 soil mesofauna, 304 excrement, 305 soil microclimate, 313 soil microfauna, 304 excrement, 304–305 soil micromorphology, 207–208 soil moisture and rhizosphere sampling, 295, 297 soil monoliths, 145 and earthworms, 310, 323 soil organic matter, 77–91 and aggregate stability, 204–207 biological fractionation, 89–91 carbon isotope identification, 90–91

Index

soil

soil

soil soil soil

soil soil

soil

soil soil

chemical fractionation, 88–89 dynamics, 311–312 formation, 84–85 functional pools, 80–82, 84 and improved soil structure, 193 labile, 83–84, 85, 88–89 mineralization, 103, 104, 106–109 modelling dynamics, 81–83 particulate matter, 87 physical fractionation, 86–88 requirements, 85 pore size distribution, 200–204 mercury injection method, 202–204, 205 water desorption method, 202, 205 porosity, 236 and macroinvertebrates, 310 methods, 198–200 processes and macroinvertebrates, 309–311 respiration measurements, 89–90 sampling design, 53–61 collection from ridging, 58 composite samples and bulking, 53–54 non-uniform areas, 55–58 plot comparisons, 58–59 random sampling, 54 sampling depth, 60 single tree effect, 59 stratification, 56–57 systematic sampling, 54 timing and frequency, 60–61 uniform areas, 54 solution composition methods, 158–163 structure, 191–208 degradation, 191–192 image analysis, 207–208 improved by trees, 192–195 and root exudates, 291 type and erosion research, 327–329 and nutrient leaching, 151–152 water, 209–234 retention, 221–222 water content methods

435

frequency domain reflectometry, 219, 220 gamma ray attenuation, 219 gravimetric, 215–216, 220 ground-penetrating radar, 219 neutron scattering, 216–217, 220 time domain reflectometry, 217–219, 220 soil water potential methods, 221–222 granular matrix sensors, 225 gypsum block sensors, 224–225 tensiometers, 222–224 soil water use methods, 232–234 Sorghum bicolor, 154, 246, 279 Sorghum sudanese, 279 Spain, 177 Sri Lanka, 177, 334 statistical analysis software, 65–66 strontium, 127, 163, 178–179 Sudan parkland water use, 214 sulphate-S, 121–122, 123 sulphur, atmospheric, and plant uptake, 125 sulphur in soil, 78 analysis by chemical fractionation, 123 analysis by physical fractionation, 123–124 deficiency problems, 121 isotopic detection techniques, 124–125 microbial, 124 mineralization from soil organic matter, 122–123 susceptibility to leaching, 152–153 Sumatra, 156–157 surface runoff, 96, 328, 329, 334 agroforestry, 224 irrigation, 211 sustainability of agricultural systems, 25, 303 swamps, 211 Swietenia macrophylla, 315

Taiwan, 177 Tamarindus indica, 277 tannins, 136

436

Tanzania, 269 taungya systems, 27 Terminalia superba, 126 termites, 161 classification, 309 excreta, 314 and organic matter availability, 314–315, 317 sphere of influence, 305, 306–307 structures, 310, 311, 322 Thailand, 16, 329 Theobroma cacao, 177, 192 Theobroma grandiflorum, 157, 175, 313, 315 tillage, 248 and destruction of soil fauna, 303 and nitrogen mineralization, 106 and soil macroinvertebrates, 317 and soil microinvertebrates, 312–313 and soil organic matter, 80 and tree root destruction, 103, 133, 237 titanium measurement, 187 Tithonia diversifolia, 126–127, 291 Togo, 85, 224 tree root growth, 173 tracer methods for leaching, 163–166 dyes, 165–166 tree canopies and erosion, 326 tree–crop competition, 99, 100, 132, 156–157, 169 and root systems, 235–236 tree–crop interaction, 8, 167, 239 complementary, 99, 101 facilitative, 99, 101 and nutrient efficiency, 98–101 and water-use efficiency, 210–211 tree fallow experiments fallow length, 49–51 fertility recovery, 49 and litter trap distribution, 141 sampling, 55 species choice, 49 tree fallows, 2, 20, 39, 45, 48–52 and crop rotation, 102 fertility and biomass production, 132

Index

fertility decline following, 50–52 and improved soil structure, 192 and nitrogen mineralization, 106, 110 and nutrient leaching, 157–158 and nutrient release, 167–168, 246–247 and root growth, 170 rotations and nutrient losses, 96 and soil organic matter, 79 and soil structure improvement, 236 tree prunings as fodder, 140 and mobile nutrients, 133 and nutrient release, 138, 168, 169 and root growth, 171 tree root biomass, 79–80 tree root depth, 169–171 and nutrient availability, 95–96, 99, 152 and nutrient capture, 167, 169 and tree–crop competition, 156–157 tree root distribution methods horizontal distribution, 244–245 profile wall, 242–244 soil coring, 240–242, 244 tree root dynamics fine root production and turnover, 249–252 ingrowth core method, 253–254 minirhizotrons, 255–257 and nutrient fluxes, 257 and production methods, 246–248 sequential excavation, 253 sequential soil coring, 248–249 tree root exudates, 296 and microbial interaction, 294 and soil structure, 291 tree root pruning, 237 tree root senescence and nitrogen transfer, 277 tree root systems, 55–56, 101, 235–257 and aluminium toxicity, 129 decomposition monitoring, 145–146, 147

Index

and experimental plots, 44–45 exposure by erosion, 341, 342 horizontal growth, 173–174 and improved soil structure, 193–194, 236 lateral, and nearby plots, 224 and nitrogen mineralization measurement, 107, 108 and nutrient capture, 167 and nutrient deficiency, 93 and nutrient efficiency, 94 and phosphorus availability, 120 pruning, 45 and radioisotope experimentation, 127 releasing carbon and nutrients, 132–133 research needs, 239 senescence and nutrient release, 168 and soil drying, 313 in soil organic matter, 77, 79, 83 sourcing nutrients, 131–132 species differentiation, 245–246 and tracing nutrient uptake, 174–179 and tree pruning, 103 and tree species selection, 236–237 and water redistribution, 213 tree root turnover, 247–248 tree species and biomass, 133 and macroinvertebrate communities, 315 selection, 263–237 trees and land disputes, 19 relationship with agriculture, 1–2 and soil fertility, 6–9 and soil structure improvement, 192–195 and soil transport rates, 325–326 spacing decisions, 237, 238 species selection for root systems, 236–237 transpiration efficiency, 214–215 and water balance, 211–214

437

Tropical Soil Biology and Fertility Programme, 303, 318

Uganda, 177 UK, 177 Ultisols, 125, 127, 156, 169 ultrasound for soil dispersion, 87–88 urea application and leaching, 154, 155 ureide production, 269–270 USA farm subsidies and agroforestry, 30

Vertisols, 151 volcanic soils, 169, 236

water penetration, 193, 207, 210 and leaching, 151–152 water pollution, 25, 106–107, 171 water resources, 14, 15 and agroforestry, 25 global trends, 210 water retention, 333 and soil organic matter, 79 water status and mycorrhizas, 274–275 water uptake and leaching, 165 and reduced leaching, 156 wildlife habitats as on-farm benefit, 26 windbreaks, 234

xylem water measurement, 232–234

Zambia fallows and mineral nitrogen, 106 intercropping and water use, 219 nitrogen mineralization measurement, 108 physical fractionation of organic matter, 110 Zimbabwe, savanna soil, 107 Zizyphus mauritiana, 277

Trees, Crops and Soil Fertility: Concepts and Research Methods

Trees, Crops and Soil Fertility Concepts and Research Methods

Trees, Crops and Soil Fertility: Concepts and Research Methods